descriptions of new dogfishes · in the follow ng ser es of papers, a rap d taxonom c approach s...

�Descriptions of new dogfishes

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Publ�shed by:CSIRO Mar�ne and Atmospher�c ResearchGPO Box 1538HobartTAS 7001AUSTRALIA

The National Library of Australia Cataloguing-in-Publication entry

Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). B�bl�ography. ISBN 1 921232 40 4 (pr�nted vers�on) ISBN 1 921232 41 2 (pdf vers�on) 1. Squalus - Identification. I. Last, P.R. (Peter Robert). II. Wh�te, W.T. (W�ll�am Toby). III. Pogonosk�, J.J (John James). IV. CSIRO. Mar�ne and Atmospher�c Research.

(Ser�es: CSIRO Mar�ne and Atmospher�c Research Paper; 14). 597.36

ISSN 1833-2331

Important Not�ce

© Copyright Commonwealth Scientific and Industrial Research Organisation (CSIRO) Australia 2007

All r�ghts are reserved and no part of th�s publ�cat�on covered by copyr�ght may be reproduced or cop�ed �n any form or by any means except w�th the wr�tten perm�ss�on of CSIRO.The results and analyses conta�ned �n th�s Report are based on a number of techn�cal, circumstantial or otherwise specified assumptions and parameters. The user must make its own assessment of the su�tab�l�ty for �ts use of the �nformat�on or mater�al conta�ned �n or generated from the Report. To the extent perm�tted by law, CSIRO excludes all l�ab�l�ty to any party for expenses, losses, damages and costs ar�s�ng d�rectly or �nd�rectly from us�ng th�s Report.

Cover Des�gn: Lou�se Bell

Layout: W�ll�am Wh�te

��Descriptions of new dogfishes

Contents

Part 1 — Appl�cat�on of a rap�d taxonom�c approach to the genus Squalus ...................................... 1

Part 2 — Squalus crassispinus sp. nov., a new spurdog of the ‘megalops-cubens�s group’ from the eastern Ind�an Ocean ........................................................................................................ 11

Part 3 — Squalus bucephalus sp. nov., a new short-snout spurdog from New Caledon�a .............. 23

Part 4 — Squalus raoulensis sp. nov., a new spurdog of the ‘megalops-cubens�s group’ from the Kermadec R�dge ................................................................................................................ 31

Part 5 — New spec�es of Squalus of the ‘highfin megalops group’ from the Australas�an reg�on ... 39

Part 6 — Descr�pt�on of Squalus chloroculus sp. nov., a new spurdog from southern Austral�a, and the resurrect�on of S. montalbani Wh�tley ........................................................................ 55

Part 7 — Two new spec�es of Squalus of the ‘m�tsukur�� group’ from the Indo–Pacific ................. 71

Part 8 — Squalus nasutus sp. nov., a new long-snout spurdog of the ‘japon�cus group’ from the Ind�an Ocean ..................................................................................................................... 83

Part 9 — Redescription of the Northern Spiny Dogfish Squalus griffini Ph�ll�pps, 1931 from New Zealand ............................................................................................................................. 91

Part 10 — Squalus hemipinnis sp. nov, a new short-snout spurdog from eastern Indones�a ......... 101

Part 11 — Clarification of the status of Squalus tasmaniensis and a d�agnos�s of Squalus acanthias from Austral�a, �nclud�ng a key to the Indo–Australas�an spec�es of Squalus .............. 109

Part 12 — DNA barcod�ng d�scr�m�nates spurdogs of the genus Squalus ..................................... 117

1Descriptions of new dogfishes

Part 1 — Application of a rapid taxonomic approach to the genus Squalus

Peter R. Last, W�ll�am T. Wh�te, John J. Pogonosk�, Dan�el C. Gledh�ll, Gordon K. Yearsley and Robert D. Ward

CSIRO Mar�ne & Atmospher�c Research, GPO Box 1538, Hobart, TAS, 7001, AUSTRALIA

INTRODUCTION

Members of the squal�d genus Squalus Bla�nv�lle, 1816 (type spec�es Squalus acanthias L�nnaeus, 1758, by subsequent des�gnat�on of G�ll, 1862), otherw�se known as spurdogs, dogsharks and dogfishes, are one of the most taxonom�cally problemat�c shark groups. Presently, 14 spec�es of Squalus are recogn�sed as val�d (Compagno et al., 2005): S. acanthias L�nnaeus, 1758; S. blainvillei (R�sso, 1827); S. cubensis Howell R�vero, 1936; S. japonicus Ish�kawa, 1908; S. megalops (Macleay, 1881); S. melanurus Fourmano�r and R�vaton, 1979; S. mitsukurii Jordan and Snyder, �n Jordan and Fowler, 1903; S. rancureli Fourmano�r and R�vaton, 1979; and 6 undescr�bed spec�es from Austral�a (Last and Stevens, 1994). At least two other nom�nal spec�es, S. brevirostris Tanaka, 1917 and S. lalannei Baranes, 2003, also appear to be val�d taxa. The status of S. acutirostris Chu, Meng and L�, 1984 and S. probatovi Myagkov and Kondyur�n, 1986 need to be determ�ned. Squalus asper Merrett, 1973 has been placed �n a related genus, Cirrhigaleus.

Squalus spec�es have been d�v�ded �nto four assemblages based on the�r morphology: the ‘acanth�as group’, the ‘bla�nv�lle� group’, the ‘megalops-cubens�s group’, and the ‘asper-barb�fer group’ (B�gelow and Schroeder, 1957; Garr�ck, 1960; Garr�ck and Paul, 1971). Members of the ‘asper-barb�fer group’ are now ass�gned to the genus Cirrhigaleus (Compagno et al., 2005). Other subgroups, whose members are cons�dered to res�de �n the ‘bla�nv�lle� group’, can be identified based on the morphology of their

component spec�es. These �nclude: the slender, long-snout spurdogs (the ‘japon�cus group’), the relat�vely short-snout spurdogs with a tall first dorsal fin, white edged caudal fin, and tricuspid denticles (the ‘highfin megalops group’), and the spurdogs w�th a moderate-s�zed snout, low dorsal fins, tricuspid denticles, and a well-defined dark caudal bar (the ‘m�tsukur�� group’). These groups may or may not represent monophylet�c groups and more research �s needed to understand the�r relat�onsh�ps. Members w�th�n groups are often very s�m�lar and have been confused �n the l�terature.

In recent years, exploratory surveys of the cont�nental slopes of Australas�a and the nearby Indo–Malay Arch�pelago, as well as an �ncreas�ng reg�onal �nterest in elasmobranchs for conservation and fishery purposes, has led to the d�scovery of many new chondr�chthyan fishes. Some of these have been discussed in important regional works. Last and Stevens (1994) identified 9 Austral�an Squalus spec�es of wh�ch 6 had not yet been formally descr�bed. Four spec�es of spurdogs were collected during recent surveys of fish markets in eastern Indonesia, of which two appear to be conspecific with undescr�bed Austral�an spec�es and the other two could not be identified to species level (White et al., 2006). S�m�larly, add�t�onal undescr�bed spurdogs were collected by French, Austral�an and New Zealand surveys �n the Tasman and Coral Seas. The Indo–West Pacific region appears to be r�ch �n spec�es of Squalus w�th more than three-quarters of the known spec�es occurr�ng there.

ABSTRACT.— The Indo–West Pacific appears to be a centre of speciation for the squaloid genus Squalus. In the follow�ng ser�es of papers, a rap�d taxonom�c approach �s used to formally descr�be 11 new spec�es and red�agnose another 4 nom�nal Squalus spec�es from the reg�on. The approach focuses on del�ver�ng a good character�sat�on of each spec�es �n a m�n�mal�st sense to �ncrease the cost effect�veness of the research and shorten t�mel�nes. Strong d�agnoses are complemented by a thorough use of d�g�tal �mages to demonstrate key morpholog�cal features. Th�s approach �s useful for �dent�fy�ng new taxa and mak�ng names ava�lable qu�ckly but �s not recommended as a replacement for formal group rev�s�onary stud�es. Morphometric characters are often poorly defined so all of the measurements taken in this study are clearly expla�ned and �llustrated. Th�s approach also �ncludes the use of molecular analyses to support morpholog�cal observat�ons.

Key words. Squaloidea – Squalidae – background – new species – Indo–West Pacific – DNA barcode

PDF contact: john.pogonosk�@cs�ro.au

2

In the follow�ng ser�es of papers (presented as Parts 2–12 of th�s �ssue), 11 new spec�es are formally descr�bed and another 4 nom�nal spec�es are red�agnosed us�ng a rap�d taxonom�c approach. The bas�s of th�s mult�-author approach �s to standard�se morphometr�c methods, restr�ct the number of spec�mens exam�ned �n deta�l, prov�de only moderate length descr�pt�ons but g�ve deta�led d�agnoses, and prov�de deta�led �mages show�ng the key features of each spec�es. Muscle t�ssue samples from 8 of these spec�es, and an add�t�onal 5 reg�onal taxa, have been analysed us�ng a molecular barcod�ng method as part of a broader �nternat�onal project that a�ms to DNA barcode all fish species (FISH-BOL, see www.fishbol.org). The results of this research (Part 12) supported the findings of morpholog�cal stud�es �n Parts 2–11.

METHODS

Morphometr�c characters were selected to d�scr�m�nate new taxa and enable compar�sons to be made w�th publ�shed �nformat�on on nom�nal Squalus spec�es from other reg�ons. Methods generally followed a w�dely adopted scheme for elasmobranchs (Compagno 1984, 2001), but focused on d�rect (po�nt-to-po�nt) rather than hor�zontal measurements. Shark researchers have made unspecified use of both approaches, so data in the literature are often unrel�able for comparat�ve purposes. Also, because the way �n wh�ch some measurements are taken can vary sl�ghtly from one shark group to another, authors need to carefully define their methods for the group in quest�on. Pr�mary measurements are expla�ned �n Table 1 and �llustrated �n F�gs 1–3. Important d�mens�ons of the snout, head, predorsal, precaudal and prenar�al reg�ons were also taken hor�zontally so compar�sons between our mater�al and l�terature spec�mens could be made when d�rect measurements were unava�lable. Measurements of the first and second dorsal-fin heights and shape of the exposed portions of the first and second dorsal-fin spines follow the concept proposed by Yamakawa et al. (1986). Several new measurements, some of wh�ch were used but not fully-defined in Springer and Burgess (1985) and then adopted by Last et al. (2002), need clarification: pre-first dorsal and pre-second dorsal lengths are direct measurements taken between the t�p of the snout and the embedded anter�or edge of the respect�ve dorsal sp�ne (just forward of the exposed sp�ne and detectable by push�ng �nto the adjacent muscle t�ssue); pre-vent length �s taken from the snout t�p to the anter�or marg�n of the ma�n open�ng of the vent; sp�racle length �s the max�mum d�ameter of the open�ng (usually measured through the vertical axis); entire lengths of the first and second dorsal fins are measured from the buried anterior edge of the spine to the apex of the free rear tip of the fin; soft dorsal-fin lengths are measured horizontally from where the rear edge of the spine joins the soft part of the fin, to the free rear tip of the fin. The lower labial furrow of spurdogs is not obv�ous externally and was not measured.

Black and white markings on the caudal fin are important for d�st�ngu�sh�ng Squalus spec�es and these characters are usually embell�shed �n late embryos and early juven�les. The most �mportant mark�ngs and the�r d�spos�t�on on the fin are illustrated in Fig. 4: the ‘caudal bar’ refers to the dark marking along the posterior edge of the fin in some spec�es (�t may be subvert�cal, extend�ng dorsally along the postdorsal caudal marg�n from the caudal fork, or obl�quely, from the or�g�n of the ventral lobe to the caudal fork); a dark ‘basal mark�ng’ �s somet�mes present near the or�g�n of the ventral lobe; a dark ‘upper caudal blotch’ and/or ‘upper caudal fr�nge’ may ex�st on the upper lobe (these markings may be sharply defined or diffuse edged); a dark ‘caudal str�pe’, wh�ch occurs rarely, �s a narrow long�tud�nal str�pe extend�ng from near the or�g�n of either the upper or lower fin lobe, along the dorsal or ventral edge of the fleshy portion of the fin. Other non-specific dark markings exist in some species. Similarly, large portions of the fin can be white, notably the distal port�on of the dorsal and ventral lobes, and the poster�or margin of the fin. In adults, white areas usually become less obv�ous and dark areas become dusky or merge w�th the main fin coloration.

The rap�d approach used �n th�s project �nvolved produc�ng m�n�mal treatments of each new spec�es and restr�ct�ng the number of spec�mens measured and rad�ographed from the type ser�es. The a�m was to prov�de names and good d�agnoses for spec�es rather than to adopt a full rev�s�onary approach. We attempted to prov�de a sol�d descr�pt�on of each spec�es to enable �t to be d�st�ngu�shed from related taxa w�thout exam�n�ng every spec�men ava�lable �n collect�ons. For example, skeletal details were not included and only extreme intraspecific var�at�ons were d�scussed. Ideally, the holotype and at least 5 post-juven�le paratypes were measured �n full, and vertebral counts were obta�ned, opt�mally for 10 types. In spec�es descr�pt�ons, morphometr�c and mer�st�c values for the holotype are given first, followed in parentheses by the ranges of the paratypes.

Mer�st�cs were obta�ned separately for trunk (monospondylous), precaudal (monospondylous + diplospondylous to origin of the caudal-fin upper lobe) and caudal (centra of the caudal fin) vertebrae. Tooth row counts, which are difficult to obtain from radiographs, were taken d�rectly from spec�mens by mak�ng �nc�s�ons at the jaw angles to expose the teeth. Type spec�mens are depos�ted �n �chthyolog�cal collect�ons ma�nly �n the Indo–Pacific region. Acronyms for all repositories follow Lev�ton et al. (1985).

DNA barcod�ng of the Squalus spec�mens �nvolved the amplification and sequencing of an approximately 655 base pa�r reg�on of the m�tochondr�al gene cytochrome ox�dase 1 (coxI). The prem�se �s that each spec�es w�ll have a un�que sequence for that coxI reg�on, or a un�que cluster of closely-related sequences (Hebert et al. 2003a). Prel�m�nary results from nearly all an�mal groups showed


Table 1. Definition of the main morphometric characters taken for Squalus. Abbreviations and definitions are adapted from Compagno (2001).

Morphometric character MethodologyTL – Total length Greatest direct distance between snout tip and caudal-fin apexPCL – Precaudal length D�rect d�stance from snout t�p to or�g�n of upper caudal lobe

PD2 – Pre-second dorsal length Direct distance from snout tip to second dorsal-fin originPD1 – Pre-first dorsal length Direct distance from snout tip to first dorsal-fin originSVL – Pre-vent length D�rect d�stance from snout t�p to anter�or end of cloacaPP2 – Prepelv�c length Direct distance from snout tip to pelvic-fin origin (use finger to find origin)PP1 – Prepectoral length Direct distance from snout tip to exposed base of pectoral fin HDL – Head length Direct distance from snout tip to upper edge of the fifth gill slitPG1 – Prebranch�al length Direct distance from snout tip to upper edge of the first gill slitPSP – Presp�racular length D�rect d�stance from snout t�p to anter�or marg�n of sp�raclePOB – Preorb�tal length Direct distance from snout tip to fleshy, anterior margin of orbitPRN – Prenar�al length D�rect d�stance from snout t�p to anter�or edge of outer nostr�lPOR – Preoral length D�rect d�stance from snout t�p to upper jaw (�nclud�ng teeth)PINL – Pre-�nner nostr�l length D�rect d�stance from snout t�p to �nner edge of nostr�lINLF – Inner nostr�l-lab�al furrow space Shortest d�stance between nostr�ls and upper lab�al furrowMOW – Mouth w�dth D�stance between ap�ces of lab�al pleats (junct�on of lab�al furrows and postoral

grooves)ULA – Lab�al furrow length D�stance from apex of lab�al pleat to anter�or edge of furrowINW – Internar�al space Shortest d�stance between the two nostr�lsINO – Interorb�tal space D�stance between soft �nterorb�t �n natural state (taken at m�d-length of eye)EYL – Eye length Length of eye, not �nclud�ng eye socketEYH – Eye he�ght He�ght of eyeSPL – Sp�racle length Max�mum w�dth of open�ngGS1 – F�rst g�ll-sl�t he�ght Vertical height of first gill slit (not following profile of gill)GS5 – F�fth g�ll-sl�t he�ght Vertical height of fifth gill slit (not following profile of gill)IDS – Interdorsal space Shortest distance between first dorsal-fin insertion and second dorsal-fin originDCS – Dorsal-caudal space Shortest distance between second dorsal-fin insertion and origin of upper caudal

lobePPS – Pectoral-pelv�c space Direct distance from pectoral-fin insertion to pelvic-fin origin (taken on ventral

s�de)PCA – Pelv�c-caudal space Direct distance from pelvic-fin insertion to origin of lower caudal lobe (taken

on ventral s�de)D1L – F�rst dorsal length Distance from first dorsal-fin origin (use thumbnail to find origin) to apex of free

rear t�pD1A – F�rst dorsal anter�or marg�n Distance from first dorsal-fin origin (use thumbnail to find origin) to point of

greatest curvature of apex of finD1B – F�rst dorsal base length Distance from first dorsal-fin origin (use thumbnail to find origin) to first dorsal-fin

�nsert�onD1H – F�rst dorsal he�ght Greatest vertical height from fin base to apex of finD1I – F�rst dorsal �nner marg�n Distance from first dorsal-fin insertion to apex of free rear tipD1P – F�rst dorsal poster�or marg�n Distance from points of greates curvature of the first dorsal-fin apex and apex of

free rear t�pD1ES – F�rst dorsal sp�ne length Distance from junction of exposed portion of spine and soft part of dorsal fin to

sp�ne apexD1BS – F�rst dorsal sp�ne base w�dth Width of exposed spine at its junction with soft dorsal finD2L – Second dorsal length Distance from second dorsal-fin origin (use thumbnail to find origin) to apex of

free rear t�p

4

Morphometric character MethodologyD2A – Second dorsal anter�or marg�n Distance from second dorsal-fin origin (use thumbnail to find origin) to point of

greatest curvature of apex of finD2B – Second dorsal base length Distance from second dorsal-fin origin (use thumbnail to find origin) to first

dorsal-fin insertionD2H – Second dorsal he�ght Greatest vertical height from fin base to apex of finD2I – Second dorsal �nner marg�n Distance from second dorsal-fin insertion to apex of free rear tipD2P – Second dorsal poster�or marg�n Distance from points of greates curvature of the second dorsal-fin apex and

apex of free rear t�pD2ES – Second dorsal sp�ne length D�stance from junct�on of exposed port�on of sp�ne and soft part of dorsal �n to

sp�ne apexD2BS – Second dorsal sp�ne base w�dth Width of exposed spine at its junction with soft dorsal finP1A – Pectoral anter�or marg�n Distance from pectoral-fin origin to apex of fin (measured from ventral surface)P1I – Pectoral �nner marg�n Distance from pectoral-fin insertion to apex of free rear tip (measured from

ventral surface)P1B – Pectoral base length Distance from pectoral-fin origin to pectoral-fin insertion (measured from

ventral surface)P1P – Pectoral poster�or marg�n Distance between points of greatest curvature of pectoral-fin apex and free rear

t�p (measured from ventral surface)P2L – Pelv�c length Distance from pelvic-fin origin (use finger to find origin) to point of greatest

curvature of apex (measured from ventral surface)P2H – Pelv�c he�ght Greatest width of pelvic fin (measured from ventral surface)P2I – Pelv�c �nner marg�n Distance from pelvic-fin insertion to apex of free rear tip (measured on ventral

surface)CDM – Dorsal caudal marg�n D�stance from or�g�n of upper caudal lobe to po�nt of greatest curvature of apex

of dorsal caudal lobeCPV – Preventral caudal marg�n D�stance from or�g�n of lower caudal lobe to po�nt of greatest curvature of apex

of ventral caudal lobeCPU – Upper postventral caudal marg�n D�stance from greatest angle of caudal fork to po�nt of greatest curvature of apex

of dorsal caudal lobeCPL – Lower postventral caudal marg�n D�stance from greatest angle of caudal fork to po�nt of greatest curvature of apex

of ventral caudal lobeCFW – Caudal fork w�dth Perpend�cular d�stance from greatest angle of caudal fork to dorsal caudal marg�nCFL – Caudal fork length D�stance from greatest angle of caudal fork to or�g�n of lower caudal lobe HANW – Head w�dth at nostr�ls W�dth of head at anter�or marg�n of nostr�ls (use stra�ght edge through anter�or

edge of nostr�ls)HAMW – Head w�dth at mouth W�dth of head at level of anter�or marg�n of mouthHDW – Head w�dth Width of head at fifth gill slitTRW – Trunk w�dth Width of body at pectoral-fin insertionsABW – Abdomen w�dth Width of body at first dorsal-fin insertionTAW – Ta�l w�dth Width of body at pelvic-fin insertionsCPW – Caudal peduncle w�dth W�dth of caudal peduncle �n front of caudal grooveHDH – Head he�ght Vertical height of head at fifth gill slitTRH – Trunk he�ght Vertical height of body at pectoral-fin insertionsABH – Abdomen he�ght Vertical height of body at first dorsal-fin insertionTAH – Ta�l he�ght Vertical height of body at pelvic-fin insertionsCPH – Caudal peduncle he�ght Vert�cal he�ght of caudal peduncle �n front of caudal grooveCLO – Clasper outer length Distance between lateral junction of pelvic-fin inner margin to apex of clasperCLI – Clasper �nner length D�stance between connect�on of the clasper base dorsally w�th the ta�l to apex of

clasper

CLB – Clasper base w�dth Width of clasper at pelvic-fin insertion

Table 1. cont’d.


HDH

TRH

ABH

TAH

CPH

HAMW

HANW

SPL

EYL

EYH

GS1

GS5

PORINW

PRN

INLF

ULA

MOW

TL (D

irec

t)

PCL

(Dir

ect)

PD2

(Dir

ect)

PD1

(Dir

ect)

A BC

Figu

re 1

. D�a

gram

mat

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enta

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qual

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vie

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. lat

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hea

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ead.

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6

D1BD1L

D1P

P1A

P1P

P1I

CDM

CPUCFW

CP

L

CPV

CFL

D1H

D1I

D1A

P2L

P2H

P2I

CLB

CLOCLI

P1B

D1ES

D1BS

D1SFL

A

B

C D

Figure 2. D�agrammat�c representat�on of the morphometr�c characters used for Squalus: A. first dorsal fin (same method used for second dorsal fin); B. pelvic fin and claspers; C. pectoral fin; and D. caudal fin. Refer to Table 1 for definitions of abbreviations and explanations of measurements.


INO

POB

PSP

HDW

TR W

ABW

TAW

CPW

PG1

HD

L

PCA

SVL

PPS

PP2

PP1

IDS

DC

S

A BA B

A B

Figu

re 3

. D�a

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pres

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hara

cter

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ex

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8

caudal stripe

upper caudalblotch

upper caudalfringe

caudal bar

white lowerlobe

basal marking

white posteriormargin

Figure 4. Terminology used for common caudal fin markings observed in Squalus.

that the chosen coxI gene reg�on �s �ndeed h�ghly spec�es-specific – exceptions include some invertebrate groups such as cn�dar�ans (Hebert et al. 2003b). Methods for barcod�ng Squalus follow Ward et al. (2005), where the barcoding of about 200 fish species showed that the technique is indeed highly effective at discriminating fish spec�es.

DISCUSSION

Prov�d�ng more effect�ve support to stakeholders (�.e. conservat�on managers, �ndustry, sc�ent�sts, and the publ�c) �s one of the major challenges for 21st Century taxonom�sts. Taxonom�c research �s often cr�t�c�sed because �t �s cons�dered to be too slow. Taxonom�sts have been tra�ned to focus on deta�l so most would prefer to complete a comprehens�ve rev�ew of a fam�ly or genus, rather than prov�de publ�shed names and s�mple d�agnoses of each spec�es. Unfortunately, th�s ph�losophy �s at odds w�th the needs of most stakeholders who s�mply w�sh to know what spec�es ex�st, the�r names, and how to distinguish them. The need for a ‘simple fix’ has �ncreased w�th the expans�on of conservat�on sc�ence and biodiversity management. Also, surveys to fill gaps in our knowledge of the b�ota on both land and �n aquat�c env�ronments, have led to the d�scovery of �ncreas�ng numbers of unidentifiable taxa, many of them new to sc�ence. Molecular approaches, such as the �nternat�onal Barcode of L�fe project campa�gn (see below), have led to the d�scovery of crypt�c spec�es and s�ster spec�es pairs, often confirming suspicions that morphological d�fferences between reg�onal morphs are more l�kely to be interspecific than population differences. In summary, the del�neat�on of operat�onal taxonom�c un�ts (OTU’s) �s �mportant but stakeholders requ�re more.

Research for the monograph Sharks and Rays of

Australia (Last and Stevens, 1994) led to the d�scovery of 97 unidentifiable taxa from the region; however less than a th�rd of these have been formally descr�bed and named s�nce �ts publ�cat�on. Some of these spec�es, such as the undescr�bed Maugean Skate (Dipturus sp. L), are now regarded as endangered and have been placed on nat�onal threatened spec�es l�sts and the IUCN Red List of Threatened Animals (IUCN, 2006). The unava�lab�l�ty of names and full descr�pt�ons of these taxa have acted as impediments to stakeholders (scientists, fishery and conservat�on managers) and have arguably, retarded progress in other fields of biological science. Increasing demands on the serv�ces of a dw�ndl�ng taxonom�c community means that faster, more efficient approaches are needed for descr�b�ng and nam�ng taxa. In th�s ser�es of papers, we have attempted to ach�eve th�s outcome by us�ng a m�n�mal�st approach but w�thout comprom�s�ng authent�c�ty and creat�ng amb�gu�ty. Electron�c publ�sh�ng enables a greater use of colour �llustrat�ons w�thout ser�ous cost �mpl�cat�ons. New electron�c journals, such as Zootaxa, have led the way by prov�d�ng h�gh qual�ty PDF’s w�th a rap�d post-subm�ss�on turn-around of manuscr�pts. In the follow�ng papers, the use of adequate descr�pt�ons and good d�agnoses helped character�se taxa and prov�ded the �nformat�on needed to d�st�ngu�sh them w�thout over-embell�shment. Skeletal �nformat�on, essent�al for systemat�c and rev�s�onary stud�es, was largely �gnored. S�m�larly, the s�zes of morphometr�c and mer�st�c datasets are kept to a m�n�mum. Us�ng th�s approach, undescr�bed taxa were treated relat�vely qu�ckly. St�ll, we stress that th�s approach w�ll never replace the need for full rev�s�onary stud�es.

We supplemented th�s approach w�th DNA barcod�ng to add a new level of taxonom�c r�gor. Newly descr�bed and the re-descr�bed spec�es had un�que coxI barcodes and these can be linked with full confidence to type material and validated identifications of these taxa. Sequences


for these spec�es have been depos�ted �n the Barcode of L�fe Database (www.boldsystems.org) and �n GenBank (www.ncbi.nlm.nih.gov). Their species-specificity and publ�c ava�lab�l�ty means that unknown spec�mens of these taxa can be correctly identified by sequencing the COI reg�on and search�ng e�ther database for a match. Only a m�nute quant�ty of t�ssue �s requ�red, wh�ch can be sent to the most conven�ent molecular laboratory for sequenc�ng. Class�cal morpholog�cal methods and molecular barcod�ng are complementary approaches that, when used together, have the potent�al to change the course of taxonomy.

ACKNOWLEDGEMENTS

A var�ety of people from var�ous �nst�tut�ons have ass�sted us in this broad taxonomic study of Indo–West Pacific spurdogs. We thank you all, the many fishers, research sc�ent�sts and research vessel crews that have helped obta�n �mportant samples from th�s reg�on. Staff from several local and �nternat�onal museums k�ndly made �mportant mater�al access�ble for parts of th�s study. Ind�v�duals that prov�ded major ass�stance to part�cular treatments are acknowledged �n appropr�ate parts of the �ssue. Three major projects, funded by Austral�an agenc�es, prov�ded mater�al and �nformat�on for most of the papers presented �n th�s �ssue. These are: a gu�de to Austral�an sharks and rays and for a study of the ecolog�cal r�sk of bycatch caught elasmobranchs (both funded by the F�sher�es Research and Development Corporat�on, FRDC) and a major study of art�sanal shark and ray fisheries of eastern Indonesia (funded by the Australian Centre for Internat�onal Agr�cultural Research, ACIAR). Another project, part�ally funded by the New Zealand Department of Conservat�on, has prov�ded support for research on New Zealand Squalus spec�es.

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and comparat�ve notes on the spec�es of th�s genus from other reg�ons. Journal of Ichthyology, 26, 1–18.

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Wh�te, W.T., Last, P.R., Stevens, J.D., Yearsley, G.K., Fahm� and Dharmad� (2006) Economically Important Sharks and Rays of Indonesia. ACIAR Monograph Ser�es, No 124, ACIAR Publ�sh�ng, Canberra, 329 pp.

Yamakawa, T., Tan�uch�, T. and Nose, Y. (1986) Rev�ew of the Etmopterus lucifer group (Squal�dae) �n Japan, p. 197–207. In: Indo–Pacific Fish Biology: Proceedings of the Second International Conference on Indo–Pacific Fishes. Tokyo. Eds: Uyeno, T., Ara�, R., Tan�uch�, T. and Matsuura, K., Ichthyolog�cal Soc�ety of Japan. Indo–Pacific Fish Biology.


Part 2 — Squalus crassispinus sp. nov., a new spurdog of the ‘megalops-cubensis group’ from the eastern Indian Ocean

Peter R. Last1, Matt Edmunds2 and Gordon K. Yearsley1

1CSIRO Mar�ne & Atmospher�c Research, GPO Box 1538, Hobart, TAS, 7001, AUSTRALIA 2Austral�an Mar�ne Ecology Pty. Ltd., 83 Parsons St, Kens�ngton, V�ctor�a 3031, AUSTRALIA

ABSTRACT.— A new spec�es of spurdog, Squalus crassispinus sp. nov., from the lower cont�nental shelf and upper cont�nental slope off Western Austral�a, �s descr�bed. It belongs to the ‘megalops-cubens�s group’, the members of which share a short snout, low dorsal fins, unicuspid denticles, and a white posterior caudal marg�n. The relat�onsh�ps of th�s spec�es to other members of the ‘megalops-cubens�s group’ are d�scussed. Squalus crassispinus can be d�st�ngu�shed from all other nom�nal Squalus spec�es by a comb�nat�on of fin coloration, dorsal-fin spine shape, denticle morphology, meristics, and morphometrics of the head, trunk and pectoral fins. Populations of S. megalops, a related, sympatr�c congener that was thought to be w�dely d�str�buted outs�de the Austral�an reg�on, are compared on an �ntrareg�onal bas�s. Characters once considered to be intraspecifically variable are found to be conservative within this region suggesting that the species may be more restricted in distribution than first thought. Squalus megalops �s re-d�agnosed based on new mater�al.

Key words. Squalo�dea – Squal�dae – Squalus crassispinus – Squalus megalops – new spec�es – Indo–West Pacific.


INTRODUCTION

Last and Stevens (1994) identified 9 Squalus spec�es from Austral�an waters of wh�ch 6 appeared to be formally undescr�bed. One of these undescr�bed spec�es belongs to the ‘megalops-cubens�s group’, wh�ch �ncludes S. megalops (Macleay, 1881), the first spurdog described from Austral�an seas. The group presently conta�ns two val�d nom�nal spec�es, S. megalops (northeastern and southeastern Atlant�c, southwestern Ind�an Ocean, western North Pacific, and Australia) and S. cubensis Howell R�vero, 1936 (western Atlant�c). Two other nom�nal spec�es, S. acutipinnis Regan, 1908 (southwest Ind�an Ocean) and S. probatovi Myagkov and Kondyur�n, 1986 (trop�cal eastern Atlant�c), are l�sted as l�kely synonyms of S. megalops (Eschmeyer, 2006). Members of the group share a short snout, low dorsal fins, unicuspid denticles, and a white posterior caudal-fin margin. In 1991, dur�ng an exploratory survey of deepwater off Western Austral�a, another member of the ‘megalops-cubens�s group’ was collected w�th S. megalops. Th�s spec�es �s formally descr�bed and an updated d�agnos�s of S. megalops �s also prov�ded.

METHODS

Methods follow those outl�ned �n Part 1 of th�s �ssue (Last

et al., 2007). The holotype (CSIRO H 2547–06) and 6 paratypes (CSIRO CA 4074, CSIRO H 1035–12, CSIRO H 1035–14, CSIRO H 1035–15, WAM P 26223–001 and WAM P 26207–005) of the new spec�es were measured �n full (Table 1). In the descr�pt�on, morphometr�c and meristic values for the holotype are given first followed �n parentheses by the ranges of the paratypes. Mer�st�cs were taken from rad�ographs of the 11 type spec�mens of the new spec�es as well as 37 spec�mens of S. megalops. Populat�ons of S. megalops from Queensland (n=3), southeastern Austral�a (n=6) and Western Austral�a (n=4), were compared morphometr�cally (Table 2). Type spec�mens are depos�ted �n the Austral�an Nat�onal F�sh Collect�on, Hobart (CSIRO), and �chthyolog�cal collect�ons of the Museum of Comparat�ve Zoology, Boston (MCZ), the Austral�a Museum, Sydney (AMS) and the Western Austral�an Museum, Perth (WAM); the�r registration numbers are prefixed with these acronyms.

Squalus crassispinus sp. nov.

F�gs 1–4; Table 1

Squalus sp. D: Last and Stevens, 1994, Sharks and Rays of Australia, pp 48, 95, figs 18, 8.34, pl. 6; Compagno, Dando and Fowler, 2005, Sharks of the World, p 80, figs, pl. 2.

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Holotype. CSIRO H 2547–06, female 580 mm TL, west of North West Cape, Western Australia, 21°37′ S, 113°59′ E, 215 m, 24 Jan 1991.Paratypes. 10 spec�mens. CSIRO CA 4074, female 497 mm TL, north of N�ckol Bay, Western Austral�a, 19° 11′ S, 116°44′ E, 194 m; CSIRO H 1035–12, female 565 mm TL, CSIRO H 1035–14, female 563 mm TL, CSIRO H 1035–15, adult male 476 mm TL, north of Dampier Archipelago, Western Australia, 19°08′ S, 116°54′ E, 196 m; CSIRO H 4649–03, female 400 mm TL, CSIRO H 4649–04, adult male 449 mm TL, north of Nickol Bay, Western Australia, 19°06′ S, 117°01′ E, 187 m; CSIRO H 4031–86, female 541 mm TL, CSIRO H 4031–87, female 543 mm TL, north of Cape Lambert, Western Australia, 18°57′ S, 117°14′ E, 248 m; WAM P 26207–005, adult male 457 mm TL, southwest of Rowley Shoals, Western Australia, 18°42′ S, 117°40′ E, 262 m; WAM P 26223–001, female 489 mm TL, north of Rowley Shoals, Western Australia, 16°32′ S, 120° 52′ E, 210 m.

DIAGNOSIS.— A small spec�es of Squalus of the ‘megalops-cubens�s group’ w�th the follow�ng comb�nat�on of characters: head w�dth at mouth 9.9–11.3% TL; mouth w�dth 2.1–2.4 t�mes hor�zontal prenar�al length; d�rect pre-second dorsal length 3.7–4.2 t�mes pectoral-fin anterior margin, 2.6–2.8 times dorsal caudal margin; preoral length 2.7–3.0 t�mes hor�zontal prenar�al length, 8.3–9.2% TL; anterior nasal flap strongly bifurcate; first dorsal fin upright to raked; exposed bases of dorsal-fin spines very broad, 1.2–1.5% TL; both dorsal fins with short inner margins, first dorsal-fin height 1.3–1.7 times its inner margin length, second dorsal-fin height 1.1–1.3 times its inner margin length; pectoral-fin anterior margin 1.9–2.2 t�mes �ts �nner marg�n length; preventral caudal margin 2.4–4.6 times inner margin of pelvic fin; caudal fin pale with poorly demarcated, whitish posterior margin, no caudal bar; flank denticles unicuspid; 39–42 (mainly 40–41) monospondylous centra, 82–86 precaudal centra, 107–111 (ma�nly 109–110) total centra; adult max�mum s�ze at least 58 cm TL.

DESCRIPTION.— Body fus�form, stout (elongate �n smallest paratypes), nape very prom�nently humped; deepest at belly (deepest at first-dorsal spine origin in smallest paratypes), max�mum depth 1.03 (0.95–1.21 �n paratypes) t�mes w�dth; trunk depth 1.01 (0.86–1.16) t�mes abdomen depth; head short 21.4 (21.0–22.3)% TL; caudal peduncle robust (less so �n smallest paratypes), elongate, 27.2 (26.6–28.4)% TL. Head not espec�ally broad, w�dth 1.03 (1.17–1.23) t�mes trunk w�dth, 1.04 (1.09–1.32) t�mes abdomen w�dth, depressed forward of spiracles, becoming subcylindrical towards pectoral-fin or�g�n, length 2.28 (2.06–2.22) �n pre-vent length; he�ght 0.84 (0.76–0.93) t�mes w�dth. Snout short, narrowly tr�angular �n lateral v�ew, apex sharply po�nted, lateral prenar�al marg�n somewhat angular (less so �n some paratypes); bluntly po�nted �n dorsal v�ew, hor�zontal length 1.32 (0.94–1.26) t�mes eye length, 0.57 (0.51–

0.66) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.70 (2.66–2.94) t�mes �n preoral length. Eye broadly oval, s�ze moderate, length 5.66 (4.53–4.89) �n head, 1.81 (1.72–2.05) t�mes he�ght; strongly notched poster�orly, weakly connected to anteroventral marg�n of sp�racle. Sp�racle moderate, crescent�c; broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.02 (2.80–3.23) �n eye length. G�ll open�ngs sl�ghtly obl�que, d�rected sl�ghtly anterodorsally from bottom to top (somet�mes upr�ght), first four subequal in size, fifth longest, height of fifth slit 2.5 (2.0–2.5)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.27 (1.11–1.27) �n preoral length; upper lab�al furrows about tw�ce length (or much more) of lower furrows, prom�nent postoral groove, subequal �n length to upper lab�al furrows (longer �n some paratypes), extend�ng posterolaterally from angle of jaws; two ser�es of funct�onal teeth �n upper jaw, 3 (2 or 3) ser�es �n lower. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Nostrils small, slightly oblique; anterior nasal flap strongly b�furcate, upper lobe largest, broadest; poster�or lobe narrow, thallate; �nternar�al space 2.21 (2.13–2.27) �n preoral length, 2.47 (2.08–2.61) t�mes nostr�l length. Dermal dent�cles (based on paratypes CSIRO H 1035–14, CSIRO H 1035–15) on flank very small, unicuspid, not �mbr�cated, crown quadrangular, w�dth subequal to length, w�th pronounced med�an r�dge, med�an r�dge commenc�ng well anter�or of rest of crown, w�th a mes�al furrow develop�ng anter�orly and converg�ng towards poster�or t�p of crown; cusp pungent, elongate; lateral extens�ons w�th a weak r�dge, cusps absent; �n paratype CSIRO H 1035–12, cusps arrowhead-shaped, broader than �n other paratypes, more densely arranged, poster�or cusp more obtuse. First dorsal fin relatively low, raked �n holotype (paratypes taller, more upr�ght), narrowly rounded ap�cally; anter�or marg�n moderately convex; upper poster�or marg�n almost stra�ght, not vert�cal, d�rected sl�ghtly anterodorsally from bottom to top (more extreme �n some paratypes), moderately concave (almost stra�ght �n CSIRO H 4649–04) near free rear t�p; free rear tip relatively thick basally, short; inner margin of fin almost stra�ght; �nsert�on of base extremely well forward of pelvic-fin origin, well posterior to free rear tip of pectoral fin; fin-spine origin above pectoral-fin insertion; sp�ne base very broad, exposed anter�orly well below junction of spine and soft portion of fin; soft portion of fin connected d�stally above m�d-po�nt of total sp�ne length; sp�ne taper�ng rap�dly d�stally, anter�or marg�n almost straight, subequal in length to second dorsal-fin spine, tip almost level with apex of soft portion of fin in undamaged types; pre-first dorsal length 3.73 (3.52–3.78) times in TL, first dorsal-fin length 1.78 (1.62–2.00) times its height, 1.21 (1.07–1.15) times second dorsal-fin length; first dorsal-fin height 1.68 (1.56–1.89) times second dorsal-fin height; exposed first dorsal spine length (0.56–0.78, holotype damaged) times height of fin. Second dorsal fin of moderate size, strongly raked; anterior margin moderately convex, apex narrowly angular, poster�or


Figure 1. Lateral v�ew of Squalus crassispinus sp. nov. holotype (CSIRO H 2547–06, female 580 mm TL).

Figure 2. Ventral v�ew of the head of Squalus crassispinus sp. nov. holotype (CSIRO H 2547–06, female 580 mm TL).

marg�n deeply concave, max�mum concav�ty sl�ghtly d�stal to m�d-po�nt of marg�n (var�able); upper port�on d�rected sl�ghtly dorsoposter�orly strongly from bottom to top; free rear t�p th�ck basally, moderately elongate, �nner margin length 0.93 (0.77–0.90) times fin height; second dorsal-fin length 2.49 (2.60–2.85) times its height; spine length 1.08 (0.96–1.11) in height of fin; fin-spine origin well behind free rear tip of pelvic fin, not exposed at level of junction with spine and fin; interdorsal space 0.94 (0.91–0.96) in length from snout tip to pectoral-fin origin, 1.16 (1.14–1.22) in pre-first dorsal length; second dorsal-fin spine with extremely broad base, subequal to width of exposed base of first dorsal-fin spine; spine slender, sharply po�nted d�stally, taper�ng very rap�dly just above

point of exposure, tip level with apex of soft portion of fin (taller than fin in some paratypes); no obvious interdorsal ridge or groove. Pectoral fin moderate, anterior margin moderately convex; �nner marg�n moderately convex, length 7.0 (6.9–7.8)% TL; apex narrowly rounded (broadly rounded �n CSIRO H 1035–12), lobe-l�ke but not falcate; poster�or marg�n moderately concave, free rear t�p bluntly angular; base very short, 2.45 (2.38–2.78) �n length of anterior margin. Pelvic fins small, anterior and poster�or marg�ns almost stra�ght, apex broadly rounded, free rear t�p angular, obtuse. Caudal peduncle long, tapering evenly to caudal fin; subcircular in cross-section anter�orly, broadly sem�c�rcular poster�orly, ventral groove well developed w�th weak med�an r�dge; lateral

14

S. crassispinus sp. nov. S. cubensis Holotype Paratypes MCZ MCZ

M�n. Max. 1462-S 1461-STL – Total length 580 457 565 282 683PCL – Precaudal length 78.6 76.1 78.9 76.2 77.9PD2 – Pre-second dorsal length 59.9 57.8 60.2 59.2 63.4PD1 – Pre-first dorsal length 26.8 26.5 28.4 27.6 28.2SVL – Pre-vent length 48.8 45.5 47.5 – 49.0PP2 – Prepelv�c length 46.7 44.0 46.0 – 49.8PP1 – Prepectoral length 21.7 21.6 22.5 22.2 21.8HDL – Head length 21.4 21.0 22.3 21.5 21.9PG1 – Prebranch�al length 17.6 17.8 18.6 18.1 18.4PSP – Presp�racular length 11.2 11.7 12.4 13.1 12.6POB – Preorb�tal length 6.9 6.9 7.3 7.2 7.6PRN – Prenar�al length 4.3 4.4 4.7 4.8 4.6POR – Preoral length 8.6 8.3 9.2 10.8 9.6MOW – Mouth w�dth 6.8 6.9 7.8 7.5 6.7ULA – Lab�al furrow length 1.8 1.8 2.2 2.2 2.1INW – Internar�al space 3.9 3.9 4.3 3.7 4.5INO – Interorb�tal space 8.8 8.6 8.8 7.9 8.3EYL – Eye length 3.8 4.5 4.9 5.5 –EYH – Eye he�ght 2.1 2.3 2.8 1.6 1.6SPL – Sp�racle length 1.2 1.4 1.7 1.7 1.9GS1 – F�rst g�ll-sl�t he�ght 2.0 1.9 2.3 1.5 1.8GS5 – F�fth g�ll-sl�t he�ght 2.5 2.0 2.5 1.5 2.1IDS – Interdorsal space 23.2 23.1 24.2 22.5 25.9DCS – Dorsal-caudal space 11.0 10.1 11.4 11.5 8.5PPS – Pectoral-pelv�c space 20.4 17.5 20.7 18.0 –PCA – Pelv�c-caudal space 27.2 26.6 28.4 22.4 23.8D1L – F�rst dorsal length 14.9 14.3 15.0 14.9 15.1D1A – F�rst dorsal anter�or marg�n 13.6 12.3 13.3 12.7 12.3D1B – F�rst dorsal base length 9.9 8.8 9.8 8.6 9.6D1H – F�rst dorsal he�ght 8.4 7.2 8.9 6.4 7.6D1I – F�rst dorsal �nner marg�n 5.0 5.0 5.5 6.1 5.9D1P – F�rst dorsal poster�or marg�n 8.4 8.5 10.7 – 9.5D1ES – F�rst dorsal sp�ne length – 4.5 5.9 2.3 –D1BS – F�rst dorsal sp�ne base w�dth 1.3 1.2 1.3 0.7 1.1D2L – Second dorsal length 12.3 12.5 13.7 12.1 11.3D2A – Second dorsal anter�or marg�n 11.4 11.7 13.0 11.3 10.5D2B – Second dorsal base length 8.4 8.9 9.1 7.0 6.5D2H – Second dorsal he�ght 5.0 4.5 5.3 3.8 3.8D2I – Second dorsal �nner marg�n 4.6 3.5 4.3 5.5 5.4D2P – Second dorsal poster�or marg�n 5.8 4.8 5.4 3.9 3.9

D2ES – Second dorsal sp�ne length 5.4 4.5 5.0 4.6 6.5

D2BS – Second dorsal sp�ne base w�dth 1.5 1.3 1.4 0.9 0.9

Table 1. Proport�onal d�mens�ons as percentages of total length for the holotype (CSIRO H 2547–06) and ranges for the 6 paratypes of Squalus crassispinus sp. nov. and two paratypes of Squalus cubensis.


S. crassispinus sp. nov. S. cubensis

Holotype Paratypes MCZ MCZM�n. Max. 1462-S 1461-S

P1A – Pectoral anter�or marg�n 15.3 14.2 15.7 14.0 15.7P1I – Pectoral �nner marg�n 7.0 6.9 7.8 9.0 11.1P1B – Pectoral base length 6.3 5.4 6.2 4.6 4.6P1P – Pectoral poster�or marg�n 11.2 9.9 11.5 8.1 11.2P2L – Pelv�c length 9.3 9.3 10.6 11.5 10.8P2H – Pelv�c he�ght 5.5 4.5 5.5 3.6 5.1P2I – Pelv�c �nner marg�n 4.4 2.5 4.8 3.2 6.1CDM – Dorsal caudal marg�n 21.2 20.9 22.3 23.0 22.1CPV – Preventral caudal marg�n 11.7 11.6 12.6 12.5 11.7CPU – Upper postventral caudal marg�n 16.9 15.3 16.9 15.4 13.9CPL – Lower postventral caudal marg�n 5.9 4.8 6.2 4.2 5.8CFW – Caudal fork w�dth 6.8 6.8 7.4 6.4 6.3CFL – Caudal fork length 8.7 9.0 10.0 11.0 9.4HANW – Head w�dth at nostr�ls 6.6 6.7 7.2 7.5 6.5HAMW – Head w�dth at mouth 10.5 9.9 11.3 11.5 11.3HDW – Head w�dth 13.1 11.8 13.3 11.3 13.3TRW – Trunk w�dth 12.8 9.1 11.3 – –ABW – Abdomen w�dth 12.6 8.3 11.8 – –TAW – Ta�l w�dth 8.4 6.7 7.8 – 6.8CPW – Caudal peduncle w�dth 3.3 2.1 3.0 2.6 2.7HDH – Head he�ght 11.0 9.8 11.7 8.9 8.8TRH – Trunk he�ght 13.1 10.1 12.9 – –ABH – Abdomen he�ght 13.0 8.9 11.7 – –TAH – Ta�l he�ght 8.4 7.3 8.2 4.8 7.0CPH – Caudal peduncle he�ght 2.5 2.4 2.8 2.6 2.5CLO – Clasper outer length – 3.7 4.1 – –CLI – Clasper �nner length – 6.9 7.1 – –CLB – Clasper base w�dth – 1.2 1.6 – –

Table 1. cont’d.

keels well developed, or�g�nat�ng poster�or to �nsert�on of second dorsal fin, terminatin g much less than an eye diameter behind caudal-fin insertion; pelvic–caudal space 0.75 (0.62–0.75) �n pectoral–pelv�c space, 0.80 (0.77–0.83) �n prepectoral length; dorsal–caudal space 2.10 (2.13–2.34) �n �nterdorsal length; dorsal caudal pit well developed, ventral caudal pit weak. Caudal fin relat�vely long, upper lobe not espec�ally broad, upper postventral marg�n undulate to moderately convex; lower lobe angular (narrowly rounded �n some); dorsal caudal marg�n 1.01 (0.94–1.05) �n head length; length of lower caudal lobe 1.81 (1.72–1.88) �n upper lobe length. Vertebral centra 110 (107–111 �n 10 paratypes, ma�nly 109–110), monospondylous 41 (39–42, ma�nly 40–41), precaudal 85 (82–86) and caudal 25 (24–27, ma�nly 25–26). Teeth �n upper jaw (of paratype, CSIRO CA 4074) 14+13=27, lower jaw 12+11=23.

COLOUR.— When fresh: un�form pale grey dorsally; much paler, wh�te on ventral surface; l�ght and dark areas poorly d�fferent�ated, pale areas forward of eye and between eye and sp�racle, prom�nent dark area below sp�racle abutt�ng h�nd marg�n of eye and extend�ng just forward of anteroventral margin of eye; fins relatively pale. Dorsal-fin spines dusky, darker grey over basal membrane; basal half of soft port�on wh�t�sh, most pronounced adjacent to fin spine; distal half pale grey with narrow, black apical margin; first dorsal-fin posterior marg�n w�th equal, weakly demarcated pale basal and darker distal portions; second dorsal-fin posterior margin pale, black apical streak confined to outer anterior margin near tip of fin spine. Caudal fin pale, posterior margin and ventral lobe wh�te; narrow dusky fr�nge along dorsal caudal margin. Pectoral fin dusky on dorsal surface, anter�or marg�n darker grey�sh, poster�or marg�n wh�te;

16

Figure 3. Cusps of the flank denticles of Squalus crassispinus sp. nov. paratype (CSIRO H 1035–12, female 565 mm TL). F�eld of v�ew w�dth 1.0 mm.

A

B

Figure 4. Lateral view of the dorsal fins of Squalus crassispinus sp. nov. paratype (CSIRO H 1035–12, female 565 mm TL) – A. first dorsal fin, B. second dorsal fin.

ventral surface uniformly pale. Pelvic fin uniformly pale dorsally and ventrally. In preservat�ve: colorat�on s�m�lar to above except pale and darker areas on dorsal fins less contrasted, except for basal part of soft portion of fins; dark streak str�pe extend�ng obl�quely from caudal p�t along dorsal fleshy portion of caudal fin; dark blotch present on lower lobe of caudal fin about a spiracle width behind fin insertion. In preservative (based on female paratype CSIRO H 4649–03): s�m�lar to holotype, border between pale and dark tonal areas extend�ng from snout, near ventral marg�n of eye, above g�ll sl�ts, then d�rected towards ventral margin, united near origin of pectoral fin;

dorsal and caudal-fin coloration similar, distinct stripe extending above fleshy portion of caudal fin; anterior margin of upper caudal-fin lobe black, lobe with a median central blotch; lower lobe of caudal fin mostly uniformly pale, ev�dence of a darker basal mark�ng.

SIZE.— Females atta�n at least 580 mm TL; males probably smaller, 3 adult male paratypes 449–476 mm TL.

DISTRIBUTION.— Endem�c to the lower cont�nental shelf and upper cont�nental slope off northwestern Austral�a. Known from North West Cape (ca. 21°S, 114°E) to north of Rowley Shoals (ca. 16°30′ S, 121° E) �n 187–262 m depth, but probably occurs more w�dely �n the eastern Ind�an Ocean.

ETYMOLOGY.— The ep�thet crassispinus, wh�ch �s a comb�nat�on of the Lat�n crassus (mean�ng fat or stout) and spinus (thorn), refers to the unusually stout dorsal-fin sp�nes.

VERNACULAR.— Fatsp�ne Spurdog.

REMARKS.— Squalus crassispinus �s sympatr�c w�th �ts closest reg�onal relat�ve, S. megalops, off Western Austral�a. Both spec�es are small w�th a short snout, low dorsal fins, unicuspid denticles, and the posterior margin of their caudal fins are mostly white. However, they d�ffer ma�nly �n the relat�ve s�ze and shape of the dorsal fins and their associated spines, and their pectoral fin shapes. Squalus crassispinus has a more upr�ght first dorsal fin, with very robust fin spines (exposed base of first dorsal-fin spine 1.2–1.3 vs. 0.6–0.8% TL �n S. megalops; exposed base of second dorsal-fin spine 1.3–1.5 vs. 0.7–0.9% TL), taller fins with short inner margins (first dorsal-fin height 1.32–1.70 vs. 1.00–1.33 times its inner margin length, second dorsal-fin height 1.08–1.30 vs. 0.71–1.04 t�mes �ts �nner marg�n length), and pectoral-fin anterior margin 1.92–2.19 (vs. 1.41–1.88) t�mes �ts �nner marg�n length. It also var�es from S. megalops �n the mouth w�dth 2.12–2.36 (vs. 2.58–3.40) t�mes hor�zontal prenar�al length; d�rect pre-second dorsal length 3.67–4.21 (vs. 4.06–4.97) times pectoral-fin anter�or marg�n, 2.59–2.84 (vs. 2.84–3.24) t�mes dorsal caudal marg�n; preoral length 2.66–2.94 (vs. 2.93–3.84) t�mes hor�zontal prenar�al length, 8.3–9.2 (vs. 9.3–9.9)% TL; preventral caudal marg�n 2.44–4.55 (vs. 1.56–2.53) times inner margin of pelvic fin. Squalus crassispinus also has sl�ghtly more vertebrae: 39–42 (ma�nly 40–41) vs. 37–40 (ma�nly 39–40) monospondylous centra; 82–86 vs. 78–84 precaudal centra; and 107–111 (ma�nly 109–110) vs. 102–110 (ma�nly 105–106) total centra. The colorat�on of both spec�es �s s�m�lar (rather pale dorsally and ventrally) although some southern Austral�an spec�es are much darker dorsally than ventrally. The poster�or margin of the caudal fin in S. crassispinus �s pale, but unl�ke S. megalops, �t �s not sharply demarcated from the rest of the fin, nor does it have a dusky bar through the ventral lobe.


Squalus megalops (Macleay, 1881)

F�gs 5–9, Table 2

Acanthias megalops Macleay, 1881. Proc. Linn. Soc. NSW, 6(2); 367 [1882]. Holotype AMS I 16255–001, Port Jackson, New South Wales, Austral�a.

Material examined. CSIRO H 1310–03, adult male 342 mm TL, south of Saumarez Reef, Queensland, 22°35′ S, 153°40′ E, 319 m; CSIRO H 1345–01, female 331 mm TL, CSIRO H 1345–02, adult male 386 mm TL, southeast of Swain Reef, Queensland, 22°53′ S, 152° 59′ E, 325 m; CSIRO H 2688–03, adult male 400 mm TL, CSIRO H 2688–04, female 375 mm TL, CSIRO H 2688–05, �mmature male 312 mm TL, east of Coffs Harbour, New South Wales, 30°24′ S, 153°23′ E, 148 m; CSIRO C 3931, female 496 mm TL, CSIRO C 3932, female 471 mm TL, east of Laur�eton, New South Wales, ca. 31°40′ S, 152°52′ E; AMS I 16255–001 (holotype), female 565 mm TL, Port Jackson, New South Wales,

A

B

C

Figure 5. Lateral v�ew of: A. Squalus megalops from V�ctor�a (CSIRO H 3762–01, female 511 mm TL); B. S. megalops from Queensland (CSIRO H 1310–03, adult male 342 mm TL); C. Squalus megalops holotype (AMS I 16255–001, female 565 mm TL).

ca. 33°50′ S, 151°15′ E; CSIRO H 3762–01, female 511 mm TL, south of Lakes Entrance, V�ctor�a, 38° 30′ S, 148°25′ E, 220 m; CSIRO H 624–02, adult male 417 mm TL, southeast of Eddystone Po�nt, Tasman�a, 41°04′ S, 148°24′ E, 80 m; CSIRO H 6449–01, immature male 245 mm TL, off Tr�al Harbour, Tasman�a, ca. 42° S, 145° E; CSIRO H 1264–15, female 547 mm TL, CSIRO H 1264–16, female 520 mm TL, CSIRO H 1264–17, female 529 mm TL, north of Maria Island, Tasmania, 42°33′ S, 148°15′ E, 82 m; CSIRO T 1515, female 474 mm TL, CSIRO T 1516, female 442 mm TL, southeast of Mar�a Island, Tasmania, 42°49′ S, 148°19′ E, 118 m; CSIRO H 1403–01, female 494 mm TL, southeast Tasman�a, 110 m; CSIRO H 2225–01, adult male 450 mm TL, CSIRO H 6482–01, adult male 415 mm TL, CSIRO H 6482–02, adult male 429 mm TL, Storm Bay, Tasman�a, ca. 43° S, 147°30′ E; CSIRO H 5737–02, female 482 mm TL, CSIRO H 5737–03, adolescent male 405 mm TL, west of Port Davey, Tasmania, 43°20′ S, 145°35′ E, 155 m; CSIRO CA 3287, female 537 mm TL, southeast of Po�nt Culver, Great Austral�an B�ght, Western Austral�a,

18

33°47′ S, 125°24′ E, 110 m; CSIRO H 2362–01, adult male 437 mm TL, CSIRO H 2365–01, female 507 mm TL, northwest of Cape Natural�st, Western Austral�a, 33°20′ S, 114°30′ E, 435 m; WAM P 27739–001 (1 of 2), adult female 540 mm TL, Western Australia, 32°00′ S, 115°30′ E, 128 m; CSIRO H 2605–08, female 542 mm TL, CSIRO H 2605–09, adult male 462 mm TL, northwest of Rottnest Island, Western Australia, 31°43′ S, 114° 58′ E, 485 m; CSIRO H 2599–05, female 308 mm TL, west of Green Head, Western Australia, 29°58′ S, 114° 26′ E, 490 m; CSIRO H 2270–03, immature male 292 mm TL, CSIRO H 2270–04, female 357 mm TL, southwest of Geraldton, Western Australia, 29°14′ S, 113°52′ E, 556 m; CSIRO H 6368–01, southwest of Shark Bay, Western Australia, 27°08′ S, 112°45′ E, 414 m; CSIRO H 2587–02, �mmature male 274 mm TL, southwest of Shark Bay, Western Australia, 27°06′ S, 112°44′ E, 370 m; CSIRO H 822–17, female 258 mm TL, CSIRO H 822–18, female 200 mm TL, southwest of Shark Bay, Western Australia, 27°03′ S, 112°40′ E, 402 m; CSIRO H 2575–19, female 345 mm TL, west of Freyc�net Estuary, Western Australia, 26°42′ S, 112°33′ E, 456 m; CSIRO H 2567–09, adult male 415 mm TL, west of Dorre Island, Western Australia, 25°09′ S, 112°09′ E, 312 m; CSIRO H 2565–02, �mmature male 319 mm TL, CSIRO H 2565–11, �mmature male 343 mm TL, west of Bern�er Island, Western Australia, 24°51′ S, 112°06′ E, 468 m.

DIAGNOSIS.— A small spec�es of Squalus of the ‘megalops-cubens�s group’ w�th the follow�ng comb�nat�on of characters: mouth w�dth 2.6–3.4 t�mes hor�zontal prenar�al length; d�rect pre-second dorsal length 4.1–5.0 times pectoral-fin anterior margin, 2.8–3.2 t�mes dorsal caudal marg�n; preoral length 2.9–3.8 t�mes horizontal prenarial length; anterior nasal flap strongly bifurcate; first dorsal fin raked; exposed bases of dorsal-fin spines narrow, 0.6–0.9% TL; both dorsal fins short with long inner margins, first dorsal-fin height 1.0–1.3 times its inner margin length, second dorsal-fin height 0.7–1.0 times its inner margin length; pectoral-fin anterior marg�n 1.4–1.9 t�mes �ts �nner marg�n length; preventral

Figure 6. Cusps of the flank denticles of Squalus megalops from NSW (CSIRO C 3931, female 496 mm TL). F�eld of v�ew w�dth 0.7 mm.

A

B

Figure 7. Lateral view of the dorsal fins of Squalus megalops from V�ctor�a (CSIRO H 3762–01, female 511 mm TL) – A. first dorsal fin, B. second dorsal fin.

Figure 8. Juvenile coloration of the caudal fin of Squalus megalops from WA (CSIRO H 6368–01, female 210 mm TL).

caudal margin 1.6–2.5 times inner margin of pelvic fin; caudal fin dusky with broad, sharply demarcated, whitish posterior margin, no caudal bar; flank denticles unicuspid; 37–40 (ma�nly 39–40) monospondylous centra, 78–84 precaudal centra, 102–110 (ma�nly 105–106) total centra; adult max�mum s�ze at least 57 cm TL.

REMARKS.— Squalus megalops appears to be w�despread �n Austral�an waters, w�th records from both temperate and tropical seas. It has been confirmed from east of the Wh�tsunday Islands �n Queensland (ca. 20° S, 151° E) to the North West Shelf off Western Austral�a (ca. 19° S, 117° E), �nclud�ng off Tasman�a (ca. 43° S) where �t comes very close to the coast �n only a few metres depth. In the northern part of �ts Austral�an range �t occurs �n greater depths, to at least 580 m. The


Table 2. Proport�onal d�mens�ons as percentages of total length for Austral�an spec�mens of Squalus megalops from southeastern Austral�a, Western Austral�a and Queensland.

southeastern Austral�a Queensland Western Austral�a(n = 6) (n = 3) (n = 4)

Mean M�n. Max. Mean M�n. Max. Mean M�n. Max.TL – Total length 457 373 527 350 328 384 462 414 541PCL – Precaudal length 77.7 76.1 79.3 78.5 77.8 78.9 78.4 77.7 79.2PD2 – Pre-second dorsal length 61.6 60.6 62.3 61.2 60.5 62.4 61.4 60.2 62.1PD1 – Pre-first dorsal length 30.2 29.1 31.6 30.6 29.9 31.6 29.6 29.1 30.2SVL – Pre-vent length 48.5 47.6 50.1 46.5 46.1 47.2 47.9 45.9 50.4PP2 – Prepelv�c length 47.4 46.4 49.9 45.0 44.8 45.4 46.9 45.5 49.1PP1 – Prepectoral length 21.5 20.4 23.2 22.6 22.2 22.9 21.9 20.9 23.0HDL – Head length 21.9 21.0 23.7 22.8 22.6 23.0 22.0 21.4 23.2PG1 – Prebranch�al length 18.5 17.7 19.8 18.9 18.6 19.2 18.3 17.8 19.1PSP – Presp�racular length 12.5 11.5 13.5 12.9 12.8 13.0 12.3 12.1 12.7POB – Preorb�tal length 7.0 6.4 7.5 7.2 7.0 7.4 7.0 6.4 7.4PRN – Prenar�al length 3.9 3.7 4.1 4.3 4.2 4.4 4.2 3.9 4.4POR – Preoral length 9.1 8.6 9.9 9.7 9.3 9.9 9.2 8.9 9.7MOW – Mouth w�dth 8.1 7.8 8.6 8.3 8.0 8.5 8.2 7.8 8.6ULA – Lab�al furrow length 2.3 2.1 2.4 2.4 2.3 2.5 2.4 2.2 2.7INW – Internar�al space 4.5 4.3 4.7 4.7 4.6 4.9 4.5 4.2 4.8INO – Interorb�tal space 8.8 8.4 9.8 9.1 8.8 9.3 8.4 7.6 9.0EYL – Eye length 4.8 4.4 5.4 5.0 4.9 5.0 4.8 4.3 5.3EYH – Eye he�ght 2.2 1.9 2.6 2.3 2.1 2.5 2.5 2.3 2.9SPL – Sp�racle length 1.4 1.0 1.7 1.4 1.2 1.5 1.6 1.5 1.7GS1 – F�rst g�ll-sl�t he�ght 2.3 2.0 2.4 1.9 1.8 1.9 2.2 1.9 2.4GS5 – F�fth g�ll-sl�t he�ght 2.4 2.1 2.5 2.5 2.3 2.6 2.2 1.8 2.4IDS – Interdorsal space 24.8 24.0 25.3 24.6 23.2 25.8 25.3 23.7 26.0DCS – Dorsal-caudal space 10.4 9.5 10.9 12.2 11.5 12.7 10.7 9.9 12.0PPS – Pectoral-pelv�c space 22.3 20.9 26.1 19.1 18.0 20.3 22.6 20.5 24.6PCA – Pelv�c-caudal space 25.6 24.5 27.0 29.0 28.7 29.4 26.9 25.7 27.8D1L – F�rst dorsal length 14.4 13.8 15.1 13.3 12.7 13.7 14.0 13.3 14.9D1A – F�rst dorsal anter�or marg�n 11.5 11.1 12.4 11.1 10.3 12.2 11.5 10.8 12.2D1B – F�rst dorsal base length 8.2 7.9 8.9 7.6 7.2 8.0 8.3 7.7 8.9D1H – F�rst dorsal he�ght 7.0 6.1 7.4 6.4 6.2 6.6 7.2 7.0 7.5D1I – F�rst dorsal �nner marg�n 6.3 6.1 6.6 5.7 5.7 5.7 5.9 5.4 6.3D1P – F�rst dorsal poster�or marg�n 8.3 6.6 9.0 7.9 7.6 8.1 7.9 7.5 8.1D1ES – F�rst dorsal sp�ne length 3.0 2.4 3.3 3.0 2.9 3.2 3.3 3.0 3.4D1BS – F�rst dorsal sp�ne base w�dth 0.8 0.7 0.8 0.7 0.6 0.7 0.7 0.6 0.8D2L – Second dorsal length 12.0 11.0 12.7 12.1 11.6 12.8 12.2 11.8 12.8D2A – Second dorsal anter�or marg�n 10.1 9.4 10.6 9.8 9.6 10.3 10.5 10.1 11.0D2B – Second dorsal base length 7.1 6.4 7.5 7.2 6.9 7.6 7.5 7.1 8.2D2H – Second dorsal he�ght 4.0 3.6 4.6 3.7 3.2 4.0 3.9 3.7 4.3D2I – Second dorsal �nner marg�n 4.9 4.5 5.3 4.9 4.5 5.1 4.9 4.7 5.0D2P – Second dorsal poster�or marg�n 4.5 3.7 5.3 4.5 4.2 4.9 4.1 3.9 4.4D2ES – Second dorsal sp�ne length 4.3 3.6 5.0 4.6 4.0 5.0 4.5 4.2 4.6D2BS – Second dorsal sp�ne base w�dth 0.8 0.8 0.9 0.8 0.7 0.8 0.8 0.8 0.9

20

Table 1. cont’d

southeastern Austral�a Queensland Western Austral�a(n = 6) (n = 3) (n = 4)

Mean M�n. Max. Mean M�n. Max. Mean M�n. Max.P1A – Pectoral anter�or marg�n 14.3 13.6 14.9 12.5 12.3 12.6 14.3 13.7 15.1P1I — Pectoral �nner marg�n 8.2 7.4 9.2 8.4 7.7 8.8 9.0 8.4 9.7P1B – Pectoral base length 5.3 4.4 5.7 5.3 4.9 5.8 4.9 4.4 5.3P1P – Pectoral poster�or marg�n 11.6 10.8 12.7 10.4 9.6 10.9 11.2 10.3 12.3P2L – Pelv�c length 10.5 9.9 11.5 10.6 9.9 11.2 10.4 9.9 10.8P2H – Pelv�c he�ght 4.8 4.3 5.2 4.7 4.5 5.1 4.9 4.7 5.2P2I – Pelv�c �nner marg�n 5.5 4.2 6.8 5.8 5.1 6.8 5.9 4.8 6.6CDM – Dorsal caudal marg�n 20.9 20.0 21.4 20.1 19.3 20.9 20.6 20.2 21.1CPV – Preventral caudal marg�n 11.0 10.5 11.3 10.6 10.4 10.7 10.9 10.7 11.0CPU – Upper postventral caudal marg�n 15.9 15.0 16.6 14.6 14.1 15.3 15.2 14.5 15.9CPL – Lower postventral caudal marg�n 4.3 4.0 4.6 3.6 3.4 3.7 4.6 3.8 5.6CFW – Caudal fork w�dth 7.1 6.6 7.5 7.0 6.9 7.1 7.0 6.6 7.3CFL – Caudal fork length 9.5 8.9 10.2 9.5 9.3 9.8 9.4 9.2 9.6HANW – Head w�dth at nostr�ls 6.7 6.3 7.6 6.9 6.7 7.2 6.7 6.3 7.1HAMW – Head w�dth at mouth 11.6 11.1 12.8 12.5 12.3 12.7 11.6 10.9 12.6HDW – Head w�dth 13.8 13.4 14.3 13.0 12.8 13.2 13.7 13.0 14.7TRW – Trunk w�dth 12.1 11.2 13.2 10.8 10.3 11.7 12.2 11.3 14.5ABW – Abdomen w�dth 10.9 10.1 12.2 10.0 9.1 11.0 10.8 10.0 11.5TAW – Ta�l w�dth 6.8 6.4 7.2 7.2 6.9 7.4 7.0 6.4 7.6CPW – Caudal peduncle w�dth 3.0 2.4 3.6 3.0 2.9 3.1 3.0 2.8 3.3HDH – Head he�ght 10.5 9.2 11.5 10.6 10.1 11.4 10.2 9.3 10.9TRH – Trunk he�ght 11.3 9.5 13.4 11.0 10.4 12.0 11.8 10.5 13.1ABH – Abdomen he�ght 11.3 9.1 14.4 11.7 10.4 13.4 11.5 9.5 13.3TAH – Ta�l he�ght 7.1 6.0 7.6 7.3 7.1 7.6 7.2 6.3 7.7CPH – Caudal peduncle he�ght 2.4 2.3 2.5 2.2 2.2 2.3 2.4 2.2 2.5CLO – Clasper outer length 3.9 3.9 3.9 3.4 2.6 4.1 2.8 3.6 3.9CLI – Clasper �nner length 9.6 9.6 9.6 9.6 9.2 10.0 7.4 9.6 10.0CLB – Clasper base w�dth 1.8 1.8 1.8 1.6 1.5 1.6 1.1 1.4 1.6

type was collected from Port Jackson, New South Wales, but similar forms around the world have been identified as th�s spec�es (Compagno et al., 2005). In Austral�an seas, �ts closest relat�ve �s S. crassispinus from Western Austral�a. Squalus megalops has a more strongly raked dorsal fin and more delicate fin spines (exposed base of first dorsal-fin spine 0.6–0.8 vs. 1.2–1.3% TL in S. crassispinus; exposed base of first dorsal-fin spine 0.7–0.9 vs. 1.3–1.5% TL), shorter fins with longer inner margins (first dorsal-fin height 1.00–1.33 vs. 1.32–1.70 times its inner margin length, second dorsal-fin height 0.71–1.04 vs. 1.08–1.30 t�mes �ts �nner marg�n length), pectoral-fin anterior margin 1.41–1.88 (vs. 1.92–2.19) t�mes �ts �nner marg�n length, mouth w�dth 2.58–3.40 (vs. 2.12–2.36) t�mes hor�zontal prenar�al length; d�rect pre-second dorsal length 4.06–4.97 (vs. 3.67–4.21) times pectoral-fin anterior margin and 2.84–3.24 (vs. 2.59–2.84) t�mes dorsal caudal marg�n, preoral length

2.93–3.84 (vs. 2.66–2.94) t�mes hor�zontal prenar�al length, 9.3–9.9 (vs. 8.3–9.2)% TL; and preventral caudal marg�n 1.56–2.53 (vs. 2.44–4.55) t�mes �nner marg�n of pelvic fin. Squalus megalops also has sl�ghtly fewer vertebrae: 37–40 (ma�nly 39–40) vs. 39–42 (ma�nly 40–41) monospondylous centra; 78–84 vs. 82–86 precaudal centra; and 102–110 (ma�nly 105–106) vs. 107–111 (ma�nly 109–110) total centra.

The colorat�on of Austral�an S. megalops �s var�able. Northeastern forms resemble S. crassispinus be�ng almost un�formly pale dorsally and ventrally, whereas southeastern Austral�an forms are much darker dorsally than ventrally. In S. megalops, the pale poster�or marg�n of the caudal fin is usually sharply demarcated from the rest of the fin rather than weakly demarcated as in S. crassispinus. Juven�le S. megalops from Western Austral�a have a strong dark obl�que bar extend�ng from


the base of the lower caudal lobe to the caudal fork (F�g. 8). In larger spec�mens, th�s bar fades but �s often ev�dent as a dusky marg�n at the caudal fork. There also appears to be some population based, intraspecific variation in the extent of dark markings on the dorsal fins and the strength of demarcat�on of poster�or marg�ns on the pectoral and caudal fins. Some regional differences in morphometr�cs ex�st between populat�ons. Queensland S. megalops (n=3) have a shorter prepelv�c length (44.8–45.4% vs. 45.5–49.9% TL), pectoral–pelv�c space (18.0–20.3% vs. 20.5–26.1% TL), pectoral-fin anterior margin (12.3–12.6% vs. 13.6–15.1% TL), lower postventral

caudal marg�n (3.4–3.7% vs. 3.8–5.6% TL), and longer pelv�c caudal space (28.7–29.4% vs. 24.5–27.8% TL) than populat�ons from other reg�ons (�.e. southeastern Austral�a and Western Austral�a, n=10). In add�t�on, they d�ffer �n several other rat�os: pectoral–pelv�c space 0.63–0.69 vs. 0.74–1.04 t�mes pelv�c–caudal space, �nterdorsal space 1.83–2.17 vs. 2.15–2.51 t�mes dorsal–caudal space, and pre-second dorsal length 4.82–4.97 vs. 4.06–4.57 t�mes pectoral anter�or marg�n length. G�ven th�s var�ab�l�ty, a more comprehens�ve �nvest�gat�on of S. megalops populat�ons �s needed as more than one spec�es may be represented.

Figure 9. Ventral v�ew of the head of: A. Squalus megalops from V�ctor�a (CSIRO H 3762–01, female 511 mm TL); B. holotype (AMS I.16255–001, female 565 mm TL).

A B

A

B

Figure 10. Lateral v�ew of: A. Squalus cubensis from Santos, Braz�l (NUPEC unreg�stered, ca. 460 mm TL); B. Squalus cubensis paratype (MCZ 1462–S, �mmature male 282 mm TL).

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Figure 11. Ventral v�ew of the head of: A. S. cubensis (NUPEC unreg�stered, ca. 460 mm TL); B. S. cubensis paratype (MCZ 1461–S, female 683 mm TL).

Although S. megalops has been recorded from several local�t�es outs�de Austral�a (�.e. southern Afr�ca, eastern Atlantic and northwestern Pacific), some of these forms may prove to be non-conspecific. A larger, Atlantic member of the ‘megalops-cubens�s group’, Squalus cubensis (F�gs 10 and 11), has a relat�vely longer snout, darker dorsal-fin apices, more robust unicuspid denticles, and stronger dorsal-fin spines.

Other material.Squalus cubensis: MCZ 1461–S (paratype), female 683 mm TL, MCZ 1462–S (paratype), �mmature male 282 mm TL, off Havana, Cuba.

ACKNOWLEDGEMENTS

Th�s study was conducted �n parallel w�th a project on southern Australian dogfishes funded by the Fisheries Research and Development Trust (FRDC 1998/108) and the new species was first identified in a study of the Austral�an chondr�chthyan fauna (Last and Stevens, 1994), also funded by the same agency (FRDC). S�ncere thanks

go to Alasta�r Graham and John Pogonosk� for assembl�ng data on Austral�an mater�als, Lou�se Conboy for ed�t�ng �mages of the types and other mater�al, John Stevens for obta�n�ng tooth counts and ed�t�ng, W�ll�am Wh�te for h�s ed�tor�al adv�ce, Dan Gledh�ll for prepar�ng a l�terature summary, Tony Rees for ass�st�ng w�th preparat�on of SEMs, and T�m Founta�n for prepar�ng rad�ographs and obta�n�ng mer�st�c �nformat�on. Thanks also to Sally Reader (AMS) for prov�d�ng �mages of the holotype of Squalus megalops and to Mark McGrouther (AMS) and Barry Hutch�ns, Sue Morr�son and Glenn Moore (WAM) for prov�d�ng collect�on data for Austral�an mater�al and to Karsten Hartel (MCZ) for prov�d�ng collect�on data for Squalus cubensis. Thanks also go to Terry Walker and Ken Graham. Most mater�al of the new spec�es was collected by the FRV Southern Surveyor dur�ng the SS1/91 voyage off Western Australia. We thank all scientific participants and crew of th�s voyage for the�r ass�stance.

REFERENCES


Eschmeyer, W.N. (2006) The Catalogue of F�shes on-l�ne. (Cal�forn�a Academy of Sc�ences: San Franc�sco) Ava�lable from: http://www.calacademy.org/research/ichthyology/catalog/fishcatmain.asp (11 December, 2006).

Howell R�vero, L. (1936) Some new, rare and l�ttle-known fishes from Cuba. Proceedings of the Boston Society for Natural History, 41, 41–76.

Last, P. R. and Stevens, J. D. (1994) Sharks and Rays of Australia. CSIRO, Austral�a, 513 pp.

Last, P.R., Wh�te, W.T., Pogonosk�, J.J., Gledh�ll, D.C., Ward, B. and Yearsley, G.K. (2007) Part 1 — Appl�cat�on of a rap�d taxonom�c approach to the genus Squalus, p 1–10. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.


Myagkov, N.A. and Kondyur�n, V.V. (1986) Sp�ny dogfishes, Squalus (Squal�dae), of the Atlant�c Ocean and comparat�ve notes on the spec�es of th�s genus from other reg�ons. Journal of Ichthyology, 26, 1–18.

Regan, C.T. (1908) A collection of fishes from the coasts of Natal, Zululand and Cape Colony. Annals of the Natal Museum, 1, 241–255.

A

B


Part 3 — Squalus bucephalus sp. nov., a new short-snout spurdog from New Caledonia

Peter R. Last1, Bernard Séret2 and John J. Pogonosk�1

1CSIRO Mar�ne & Atmospher�c Research, GPO Box 1538, Hobart, TAS, 7001, AUSTRALIA 2Muséum Nat�onal d’H�sto�re Naturelle, Département Systémat�que et Evolut�on, UMS n° 602,

“Taxonom�e et Collect�ons”, case postale 51, 55 rue Buffon, 75231 Par�s cedex 05, FRANCE

ABSTRACT.— A new spec�es of spurdog, Squalus bucephalus sp. nov., �s descr�bed from deepwater south of New Caledon�a �n the northern Tasman Sea. It belongs to the ‘megalops-cubens�s group’ but d�ffers from Austral�an forms of S. megalops in having a broader head, larger dorsal-fin spines and reaches a larger adult s�ze. It also d�ffers �n several other mer�st�c and morphometr�c deta�ls and �s the only Squalus known to possess both un�cusp�d and mult�cusp�d dent�cles �n adults. It �s morpholog�cally s�m�lar to the newly descr�bed S. crassispinus from the eastern Indian Ocean, but differs in having a lower, strongly raked first dorsal fin, more vertebrae, and more slender dorsal-fin spines.

Key words. Squalo�dea – Squal�dae – Squalus bucephalus – new spec�es – New Caledon�a.


INTRODUCTION

In recent decades, the French research �nst�tute IRD (formerly ORSTOM) has conducted a ser�es of deepwater marine biodiversity surveys of the South Pacific (Richer de Forges, 1990, 1996; Séret, 1997). Some of these voyages have targeted the chondr�chthyan fauna and these collect�ons are presently under �nvest�gat�on (Séret and Last, unpubl. data). A prel�m�nary l�st of chondr�chthyan fishes published by Séret (1994), included new species. The total number of chondrichthyan fishes in the econom�c zone of New Caledon�a �s currently est�mated at 73, compr�s�ng 51 sharks, 18 rays and 4 ch�maeras (Séret, unpubl�shed data). Two endem�c spec�es of Squalus have been recorded from the adjacent reg�on. Squalus melanurus Fourmano�r and R�vaton, 1979 was descr�bed from mater�al collected on the �nsular slopes of the Uat�o and Bular� Passes near New Caledon�a, and S. rancureli Fourmano�r and R�vaton, 1979 from �nsular slopes of nearby Vanuatu. In the recent surveys of th�s reg�on, an undescr�bed, short-snout spurdog belong�ng to the ‘megalops-cubens�s group’ was collected by one of us (B. Séret). Th�s spec�es �s formally descr�bed and compared to other nom�nal members of the subgroup, �.e. S. cubensis Howell R�vero, 1936 and S. megalops (Macleay, 1881), and a new spec�es descr�bed �n th�s �ssue, S. crassispinus Last, Edmunds and Yearsley, 2007 (see Part 2).

METHODS

Methods generally follow those outl�ned �n Part 1 of th�s �ssue (Last et al., 2007b). Ava�lable mater�al cons�sted of the holotype and three paratypes (Table 1). The largest spec�mens, two adult males (NMNZ P 34030), had been ev�scerated so a smaller �mmature male �n better cond�t�on was selected as the holotype. In the descr�pt�on, morphometr�c and mer�st�c values for the holotype are given first followed in parentheses by the ranges of the measured paratypes. Measurements affected by damage to the largest paratypes were excluded so �nformat�on for only one paratype �s ava�lable �n some �nstances. Mer�st�cs were taken from rad�ographs. Type spec�mens are depos�ted �n the �chthyolog�cal collect�ons of the Muséum nat�onal d’H�sto�re naturelle, Par�s (MNHN) and Nat�onal Museum of New Zealand, Well�ngton (NMNZ); their registration numbers are prefixed with these acronyms.

Squalus bucephalus sp. nov.


Squalus sp. n.: Séret, 1994 (l�sted �n Table A, p. 7); Séret �n Grandperr�n et al., 1997 (l�sted �n Annexe 11 – L�ste des Po�sons, p. 112)

24

Figure 1. Lateral v�ew of Squalus bucephalus sp. nov. holotype (MNHN 2006–1754, �mmature male 556 mm TL).

Figure 2. Ventral v�ew of the head of Squalus bucephalus sp. nov. holotype (MNHN 2006–1754, �mmature male 556 mm TL).

Holotype. MNHN 2006–1754, �mmature male 556 mm TL, Norfolk Ridge, south of New Caledonia, 23°43′ S, 168°16′ E, 405–411 m, 28 November 1993.Paratypes. 3 spec�mens. MNHN 1997–3641, female 430 mm TL, Stylaster Seamount, Norfolk R�dge, New Caledonia, 23°37′ S, 167°42′, 420–470 m; NMNZ P 34030, two adult males 790, 808 mm TL, Stylaster Seamount, Norfolk Ridge, New Caledonia, 23°35′ S, 167°42′ E, 448–880 m.

DIAGNOSIS.— A large, broad-headed spec�es of Squalus of the ‘megalops-cubens�s group’ w�th the follow�ng comb�nat�on of characters: head w�dth at mouth 12.1–13.5% TL; mouth w�dth 2.2–2.7 t�mes

hor�zontal prenar�al length; d�rect pre-second dorsal length 3.9–4.0 times pectoral-fin anterior margin, 2.7 t�mes dorsal caudal marg�n; preoral length 2.7–3.1 t�mes hor�zontal prenar�al length, 9.2–10.5% TL; anter�or nasal flap strongly bifurcate; first dorsal fin low, raked; exposed bases of dorsal-fin spines broad, 0.7–0.8% TL; both dorsal fins with short inner margins, first dorsal-fin height 1.4–1.5 times its inner margin length, second dorsal-fin height 0.8–1.0 times its inner margin length; pectoral-fin anter�or marg�n 1.8–2.0 t�mes �ts �nner marg�n length; preventral caudal marg�n 1.8–2.2 t�mes �nner marg�n of pelvic fin; dorsal fins with dark distal portion, posterior margin of free rear tip pale; caudal fin upper posterior marg�n broadly wh�te, lower lobe w�th dark base and


pale outer half, no caudal bar; flank denticles unicuspid or weakly tr�cusp�d; 45 monospondylous centra, 86–89 precaudal centra, 113–118 total centra; adult max�mum s�ze at least 90 cm TL.

DESCRIPTION.— Body fus�form, relat�vely short, robust (small paratype more deep-bod�ed, mature males elongate), nape prominently humped; deepest near first dorsal-fin spine origin, maximum depth 0.97 (1.02 in paratype) t�mes w�dth; trunk depth 1.01 (1.04) t�mes abdomen depth; head short 21.0 (21.8–22.9)% TL; caudal peduncle length relat�vely short, 23.3 (23.4–24.5)% TL. Head very robust, broad, sl�ghtly w�der than trunk, w�dth 1.17 (1.07) t�mes trunk w�dth, 1.33 (1.29) t�mes abdomen w�dth, depressed forward of sp�racles, almost subtriangular in cross-section towards pectoral-fin or�g�n, length 2.24 (2.08–2.31) �n pre-vent length; he�ght 0.72 (0.66–0.72) t�mes w�dth. Snout short, narrowly tr�angular �n lateral v�ew, apex bluntly po�nted, lateral prenar�al marg�n not angular; very broadly rounded �n dorsal v�ew, hor�zontal length 1.29 (1.12–1.18) t�mes eye length, 0.61 (0.55–0.59) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.73 (2.78–3.07) t�mes �n preoral length. Eye narrowly oval, s�ze moderate, length 4.71 (4.72–4.93) �n head, 2.25 (2.26–2.65) t�mes he�ght; strongly notched poster�orly, notch extend�ng halfway to sp�racle as a well-developed furrow. Sp�racle small, crescent�c; very broad, bulbous lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.11 (3.27–4.31) �n eye length. Gill openings short, almost upright, first four subequal in size, fifth longest, height of fifth slit 2.1 (1.8–2.5)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.08 (1.13–1.25) �n preoral length; upper lab�al furrow much longer than lower furrow; postoral groove prom�nent, much longer than upper lab�al furrows, extend�ng posterolaterally from angle of jaws; one ser�es of funct�onal teeth �n upper jaw (1–2 �n paratypes), 1–2 ser�es �n lower jaw. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than

Figure 3. Cusps of the flank denticles of Squalus bucephalus sp. nov. paratype (NMNZ P 34030, adult male 808 mm TL). F�eld of v�ew w�dth 1.6 mm.

Figure 4. Lateral view of the dorsal fins of Squalus bucephalus sp. nov. holotype (MNHN 2006–1754, immature male 556 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B

length of cusp. Nostr�ls small, sl�ghtly obl�que; anter�or nasal flap strongly bifurcate, upper lobe largest, broadest; posterior lobe short, finger-like; internarial space 2.01 (1.75–2.11) �n preoral length, 2.36 (2.71–2.77) t�mes nostr�l length. Dermal dent�cles (based on holotype) on flank very small, not imbricated, varying in shape from weakly tr�cusp�d to un�cusp�d. Largest dent�cles tr�cusp�d, central cusp enlarged w�th strong med�an r�dge; lateral cusps short, weakly po�nted, w�thout r�dges. Smallest dent�cles extremely compressed, anv�l shaped �n lateral profile, central cusp with a narrow median ridge; bluntly po�nted d�stally; often w�th short, nodular, lateral extens�ons; �n mature male paratype NMNZ P 34030, denticles below dorsal fin mostly broadly unicuspid with long, broad central cusps and bulg�ng lateral keels (keels somet�mes expanded to form weak tr�cusp�d crowns). Other dent�cles var�able �n shape, w�th rud�mentary lateral cusps of vary�ng s�zes; ap�ces usually sharply pointed. First dorsal fin low, raked slightly, narrowly rounded ap�cally (more upr�ght �n adult male paratypes); anter�or marg�n weakly convex; upper poster�or marg�n almost stra�ght, subvert�cal, strongly concave near free rear t�p; free rear t�p rather deep basally, short; �nner margin of fin almost straight; insertion of base well forward of pelvic-fin origin, slightly posterior to free rear tip of pectoral fin; fin-spine origin above (just behind in one paratype) pectoral-fin insertion; spine base narrow,

26

exposed anter�orly near junct�on of sp�ne and soft port�on of fin; soft portion of fin connected near mid-point of total sp�ne length (d�stal port�on of sp�ne usually abraded); sp�ne taper�ng gradually d�stally, anter�or marg�n almost stra�ght, exposed port�on t�lted, shorter than length of exposed portion of second dorsal-fin spine; pre-first dorsal length 3.62 (3.47–3.54) times in TL; first dorsal-fin length 1.90 (1.75–1.93) t�mes �ts he�ght, 1.32 (1.23–1.34) times second dorsal-fin length; first dorsal-fin height 2.02 (1.69–2.22) times second dorsal-fin height; exposed first dorsal spine length 0.41 (0.32) times height of fin. Second dorsal fin small, very strongly raked; anterior margin weakly convex, apex narrowly rounded; poster�or marg�n deeply concave, max�mum concav�ty almost near m�d-po�nt of marg�n, upper port�on d�rected dorsoposter�orly strongly from bottom to top; free rear t�p elongate, �nner margin length 1.18 (0.97–1.04) times fin height; second dorsal-fin length 2.90 (2.61–2.90) times its height; spine length 1.09 (0.82) in height of fin; fin-spine origin well behind free rear tip of pelvic fin, exposed about at level of junction with spine and soft portion of fin; second spine narrow based (broader �n adult males), slender, sharply po�nted d�stally, not taper�ng rap�dly just above po�nt of exposure, sp�ne t�p extend�ng beh�nd level of �nsert�on of fin (when intact), much more strongly raked than first sp�ne; �nterdorsal space 0.93 (0.89–1.01) �n length from snout tip to pectoral-fin origin, 1.21 (1.13–1.29) in pre-first dorsal length; weak interdorsal groove. Pectoral fin moderate, anterior margin moderately convex; inner marg�n weakly convex, length 8.2 (7.4–8.2)% TL; apex narrowly rounded, lobe-l�ke but not falcate; poster�or marg�n weakly to moderately concave, free rear t�p narrowly rounded; base very short, 3.14 (2.58–2.82) �n length of anterior margin. Pelvic fins small, anterior and poster�or marg�ns almost stra�ght (somet�mes weakly convex), apex broadly rounded, free rear t�p bluntly po�nted. Caudal peduncle short, pelv�c–caudal space 23.3 (23.4–24.5)% TL; tapering evenly to caudal fin; subcircular in cross-sect�on anter�orly, broadly sem�c�rcular poster�orly, ventral groove usually very well developed; lateral keels well developed, the�r or�g�n obscure, about under insertion of second dorsal fin, terminating about half an eye diameter behind caudal-fin insertion; pelvic–caudal space 0.86 (0.84–0.93) �n pectoral–pelv�c space, 0.91 (0.93–0.94) �n prepectoral length; dorsal–caudal space 2.42 (2.24–2.72) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal pit rudimentary. Caudal fin relat�vely long, dorsal caudal marg�n 0.86 (0.98–1.00) �n head length; length of lower caudal lobe 2.00 (1.92–2.05) �n upper lobe length; upper poster�or lobe moderately convex; lower lobe apex narrowly rounded. Vertebral centra 118 (113–117 �n paratypes), monospondylous 45 (45), precaudal 88 (86–89) and caudal 30 (27–28). Teeth �n upper jaw of holotype 13+1+13 = 27, lower jaw 11(or 12)+11(or 12) = 22(or 23, 24), uncut side of jaw difficult to count, smaller symphys�al tooth w�th cusp d�rected sl�ghtly to r�ght; �n paratype, 13+13 = 26, lower jaw 11+12 = 23.

COLOUR.— In preservat�ve: dorsal half un�formly dark brown; paler yellow�sh or wh�te ventrally (paratypes w�th sl�ghtly darker, dusky tones on belly and ventral surface of ta�l, blotches on adult males); l�ght and dark tones sharply demarcated on head (tones gradat�ng on abdomen and ta�l), pale boundary extend�ng around angle of snout just below eye and through upper two th�rds of gill slits. First dorsal-fin base pale brown; outer fin dark grey to black (more pronounced �n smallest paratype, NMNZ P34030), most pronounced at fin apex; free rear t�p dusky w�th wh�t�sh poster�or marg�n; adults w�th uniform dusky fins, upper posterior margins translucent. Second dorsal-fin base slightly paler than dorsal surface of body (more obv�ous �n NMNZ P34030); outer port�on of fin dark grey (distinctly black in NMNZ P34030); free rear t�p grey�sh w�th narrow wh�t�sh marg�n extend�ng to angle of fin. Caudal fin central fleshy portion brownish, upper lobe mostly darker, grey�sh to black�sh; basal half of lower lobe w�th tr�angular dark grey to black bar; posterior margin of fin sharply demarcated pale yellow to wh�te (more extens�ve and strongly pronounced �n smallest paratype); l�ght and dark �nterface extend�ng from about m�ddle of preventral marg�n almost �n a stra�ght l�ne dorsoposter�orly towards apex of upper lobe (anteroposter�or marg�n of dorsal lobe pale �n smallest paratype); posterior angle of caudal fin lacking black bar; adult males with more uniformly coloured fin, pale posterior margin strongly demarcated. Pectoral fin upper surface dark grey (darker than dorsal colorat�on) w�th a pronounced pale poster�or marg�n; ventral surface with pale base and greyish outer fin, posterior margin pale. Pelvic fin brownish or greyish, of a similar tone or sl�ghtly darker than dorsal surface of body, poster�or marg�n pale; ventral surface pale or blotched, s�m�lar to ton�ng of belly.

SIZE.— Large spec�es known from only a few �nd�v�duals; two largest males both mature (790– 808 mm TL); smallest male (556 mm TL) had undeveloped claspers. Add�t�onal non-type spec�mens �n the MNHN

Figure 5. Caudal fin coloration of juvenile of Squalus bucephalus sp. nov. paratype (MNHN 1997–3641, female 430 mm TL).


Table 1. Proport�onal d�mens�ons as percentages of total length for the holotype (MNHN 2006–1754) and ranges for the 3 paratypes of Squalus bucephalus sp. nov.

S. bucephalus sp. nov. Holotype Paratypes

M�n. Max.TL – Total length 556 430 808PCL – Precaudal length 74.5 76.3 77.8PD2 – Pre-second dorsal length 57.6 58.8 61.1PD1 – Pre-first dorsal length 27.7 28.2 28.8SVL – Pre-vent length 47.0 47.8 50.4PP2 – Prepelv�c length 46.9 45.6 48.8PP1 – Prepectoral length 21.3 22.0 22.6HDL – Head length 21.0 21.8 22.9PG1 – Prebranch�al length 18.5 18.5 20.1PSP – Presp�racular length 13.0 12.6 13.9POB – Preorb�tal length 7.7 7.1 7.9PRN – Prenar�al length 4.7 4.6 5.2POR – Preoral length 9.2 9.2 10.5MOW – Mouth w�dth 8.5 8.1 8.4ULA – Lab�al furrow length 2.1 2.2 2.4INW – Internar�al space 4.6 5.0 5.3INO – Interorb�tal space 9.4 8.6 9.8EYL – Eye length 4.4 4.4 4.9EYH – Eye he�ght 2.0 1.8 2.0SPL – Sp�racle length 1.4 1.1 1.4GS1 – F�rst g�ll-sl�t he�ght 2.0 1.4 2.0GS5 – F�fth g�ll-sl�t he�ght 2.1 1.8 2.5IDS – Interdorsal space 22.9 22.3 25.3DCS – Dorsal-caudal space 9.5 9.3 9.9PPS – Pectoral-pelv�c space 20.0 20.0 21.7PCA – Pelv�c-caudal space 23.3 23.4 24.5D1L – F�rst dorsal length 16.1 14.8 15.7D1A – F�rst dorsal anter�or marg�n 14.2 12.5 14.6D1B – F�rst dorsal base length 10.1 9.1 10.0D1H – F�rst dorsal he�ght 8.5 8.1 8.4D1I – F�rst dorsal �nner marg�n 6.1 5.4 5.8D1P – F�rst dorsal poster�or marg�n 9.4 9.2 9.8D1ES – F�rst dorsal sp�ne length 3.5 – 2.6D1BS – F�rst dorsal sp�ne base w�dth 0.8 0.8 0.8D2L – Second dorsal length 12.2 11.0 12.7D2A – Second dorsal anter�or marg�n 10.7 9.4 11.4D2B – Second dorsal base length 7.5 7.0 7.9D2H – Second dorsal he�ght 4.2 3.8 4.8D2I – Second dorsal �nner marg�n 5.0 4.0 4.8D2P – Second dorsal poster�or marg�n 5.0 4.1 5.2D2ES – Second dorsal sp�ne length 4.6 – 3.8D2BS – Second dorsal sp�ne base w�dth 0.8 0.7 0.8

28

Table 1. cont’d

S. bucephalus sp. nov. Holotype Paratypes

M�n. Max.P1A – Pectoral anter�or marg�n 15.8 14.3 15.7P1I – Pectoral �nner marg�n 8.2 7.4 8.2P1B – Pectoral base length 5.0 5.1 5.9P1P – Pectoral poster�or marg�n 12.3 11.4 12.4P2L – Pelv�c length 9.4 9.0 11.0P2H – Pelv�c he�ght 5.0 4.6 6.0P2I – Pelv�c �nner marg�n 5.5 5.4 6.2CDM – Dorsal caudal marg�n 24.5 21.9 23.5CPV – Preventral caudal marg�n 12.3 11.4 11.5CPU – Upper postventral caudal marg�n 18.3 17.1 17.7CPL – Lower postventral caudal marg�n 5.2 4.8 5.5CFW – Caudal fork w�dth 7.1 7.1 7.3CFL – Caudal fork length 9.6 9.3 9.7HANW – Head w�dth at nostr�ls 8.8 7.7 9.3HAMW – Head w�dth at mouth 13.0 12.1 13.5HDW – Head w�dth 13.8 13.7 14.2TRW – Trunk w�dth 11.8 – 12.9ABW – Abdomen w�dth 10.3 – 10.7TAW – Ta�l w�dth 6.6 6.3 6.6CPW – Caudal peduncle w�dth 3.1 3.0 3.3HDH – Head he�ght 9.9 9.3 10.0TRH – Trunk he�ght 11.5 – 13.1ABH – Abdomen he�ght 11.3 – 12.6TAH – Ta�l he�ght 6.9 6.6 6.9CPH – Caudal peduncle he�ght 2.7 2.6 2.8CLO – Clasper outer length 1.0 2.5 2.9CLI – Clasper �nner length 3.2 8.7 8.8CLB – Clasper base w�dth 0.8 1.7 1.7

ranged �n s�ze from 301–900 mm TL; smallest adult male 662 mm TL.

DISTRIBUTION.— Known only from the northern Tasman Sea on the Norfolk R�dge (Stylaster Seamount), and off New Caledon�a, �n 448–880 m depth.

ETYMOLOGY.— Der�ved from the Lat�n bu (large) and Greek kephalis (of the head) �n allus�on to �ts relat�vely bulky head.

VERNACULAR.— B�ghead Spurdog (Engl�sh), A�gu�llat à grosse tête (French).

REMARKS.— Squalus bucephalus can be d�st�ngu�shed from all other members of the ‘megalops-cubens�s group’

by the comb�nat�on of an unusually broad head, large dorsal-fin spines, and other meristic and morphometric deta�ls. It �s the only Squalus known to possess both un�cusp�d and mult�cusp�d dent�cles �n adults. It d�ffers from S. megalops �n reach�ng a larger s�ze (males about 800 vs. 470 mm TL �n S. megalops), having different first dorsal fin (first dorsal-fin height 1.4–1.5 vs. 1.0–1.3 times �ts �nner marg�n length) and preoral shapes (mouth w�dth 2.2–2.7 vs. 2.6–3.4 t�mes hor�zontal prenar�al length), and more vertebrae (45 vs. 37–40 monospondylous centra, 86–89 vs. 78–84, and rarely more than 82, precaudal centra, and 113–119 vs. 102–110, and rarely more than 106, total centra). It �s morpholog�cally closer to S. crassispinus but differs in having a lower, more strongly raked first dorsal fin, broader head (width at mouth 12.1–13.5% vs. 9.9–11.3% TL �n S. crassispinus), longer preoral length


(9.2–10.5% vs. 8.3–9.2% TL), less robust exposed bases of dorsal-fin spines (0.7–0.8% vs. 1.2–1.5% TL), a better demarcated, pale posterior caudal-fin margin, and more vertebrae (45 vs. 39–42 monospondylous centra, 86–89 vs. 82–86 precaudal centra, 113–119 vs. 107–111 total centra). The second dorsal-fin height is 0.9–1.0 (vs. 1.1–1.3) t�mes �ts �nner marg�n length and the preventral caudal marg�n �s 1.8–2.2 (vs. 2.4–4.6) t�mes the length of the inner margin of the pelvic fin. Another member of the ‘megalops-cubens�s group’, S. cubensis from the Atlant�c Ocean has a longer, narrower snout and darker dorsal fins (see Last et al., 2007a, Part 2 of th�s �ssue).

ACKNOWLEDGEMENTS

The authors w�sh to thank Alasta�r Graham for obta�n�ng local�ty data, Adr�an Turner (Un�vers�ty of Auckland) for prepar�ng the SEMs, Andrew Stewart (NMNZ), Cather�ne Hobb�s (Un�vers�ty of Auckland) and Lou�se Conboy (CSIRO) for obta�n�ng �mages of the types, John Stevens for obta�n�ng tooth counts and ed�t�ng, Sp�key R�ddoch for prepar�ng rad�ographs, Dan�el Gledh�ll for prepar�ng a l�terature summary, and W�ll�am Wh�te and Gordon Yearsley for the�r ed�tor�al comments. Thanks also to Bertrand R�cher de Forges, leader of the IRD/MNHN exploratory cru�se program off New Caledon�a and René Grandperr�n, ch�ef sc�ent�st of the HALIPRO cru�se.

REFERENCES

Fourmano�r, P. and R�vaton, J. (1979) Po�ssons de la pente réc�fale externe de Nouvelle-Calédon�e et des Nouvelles-Hébr�des. Cahiers de l’Indo-Pacifique, 1, 405–443.

Grandperr�n, R., Farman, R., Lorance, P., Jomessy T., Hamel P., Laboute P., Labrosse, P., R�cher de Forges, B., Séret, B. and V�rly, S. (1997) Campagne HALIPRO 2 de chalutage exploratoires profonds dans le sud de la zone économique de Nouvelle-Calédonie (R.V. Tangaroa, 4-28 novembre 1996). ORSTOM, Nouméa, Rapport de m�ss�on, 150 pp.

Howell R�vero, L. (1936) Some new, rare and l�ttle-known fishes from Cuba. Proceedings of the Boston Society of Natural History. 41, 41–76.

Last, P.R. and Stevens, J.D. (1994) Sharks and Rays of Australia. CSIRO Publ�cat�ons, Melbourne, 513 pp.

Last, P.R., Edmunds, M. and Yearsley, G.K. (2007a) Part 2 — Squalus crassispinus sp. nov., a new spurdog of the ‘megalops-cubens�s group’ from the eastern Ind�an Ocean, p. 11–22. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Last, P.R., Wh�te, W.T., Pogonosk�, J.J., Gledh�ll, D.C., Ward, B. and Yearsley, G.K. (2007b) Part 1 — Appl�cat�on of a rap�d taxonom�c approach to the genus Squalus, p. 1–10. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Macleay, W. (1881) Descr�pt�ve catalogue of Austral�an fishes. Part IV. Proceedings of the Linnean Society of New South Wales. 6, 202–387.

R�cher de Forges, B. (1990) Les campagnes bathyales dans la zone économ�que de la Nouvelle-Calédon�e. In: Résultats des Campagnes MUSORSTOM, Vol. 6. Ed: Crosn�er, A., Mémoires du Muséum National d’histoire Naturelle, Série A, 145, 9–54.

R�cher de Forges, B. (1996) Les campagnes d’échant�llonnage du benthos bathyal en Nouvelle-Calédon�e, en 1993 et 1994 (BATHUS 1 à 4, SMIB 8, HALIPRO 1). In: Résultats des Campagnes MUSORSTOM, Vol. 15. Ed: Crosn�er, A., Mémoires du Muséum National d’histoire Naturelle, Série A, 168, 9–54.

Séret, B. (1994) Chondrichthyan fishes of New Caledonia. Chondros, 5(3), 6–9.

Séret, B. (1997) Po�ssons de profondeur de Nouvelle-Calédon�e: apports des campagnes MUSORSTOM. Deep water fishes of New-Caledonia: contributions of the MUSORSTOM cru�ses. In: Séret, B. (ed.), Résultats des campagnes MUSORSTOM, vol. 17. Mémoires du Muséum National d’histoire Naturelle, Série A, 174, 9–16.


Part 4 — Squalus raoulensis sp. nov., a new spurdog of the ‘megalops-cubensis group’ from the Kermadec Ridge

Cl�nton A.J. Duffy1 and Peter R. Last2

1Department of Conservat�on, Pr�vate Bag 68908 Newton, Auckland 1145, NEW ZEALAND2CSIRO Mar�ne & Atmospher�c Research, GPO Box 1538, Hobart, TAS, 7001, AUSTRALIA

ABSTRACT.— A new spec�es of spurdog, Squalus raoulensis sp. nov., �s descr�bed from the upper �nsular slope of Raoul Island, Kermadec Islands, and the Kermadec R�dge north of Raoul Island, New Zealand. The new spec�es belongs to a subgroup of Squalus whose members share a relat�vely short snout and un�cusp�d dent�cles. Squalus raoulensis differs from its closest congeners primarily in a combination of fin coloration, dorsal-fin shape, meristics, and morphometrics of the head, trunk and pectoral fins. The relationships of this spec�es to other members of th�s subgroup are d�scussed.

Key words. Squalo�dea – Squal�dae – Squalus raoulensis – new species – south-west Pacific – Kermadec R�dge.


INTRODUCTION

Squalus acanthias L. and S. griffini Ph�ll�pps, 1931 are the only Squalus spec�es currently recogn�sed from New Zealand waters (Garr�ck, 1960; Paul�n et al., 1989; Duffy and Last, 2007, Part 9 th�s �ssue). Squalus acanthias �s w�dely d�str�buted around the North and South Islands, Chatham R�se and Campbell Plateau, whereas S. griffini �s restr�cted to waters north of the Subtrop�cal Front. Dur�ng an exam�nat�on of Squalus held in the fish collect�on of the Museum of New Zealand, a th�rd New Zealand species was identified based on a single female specimen collected by a scientific observer from the Kermadec R�dge, north of Raoul Island. Subsequently, the sen�or author collected two mature male spec�mens from the upper �nsular slope off Raoul Island �n the Kermadec group. These spec�mens are descr�bed here�n as a new spec�es belong�ng to the ‘megalops-cubens�s group’, and compar�sons are made w�th s�m�lar spec�es, S. bucephalus Last, Seret and Pogonosk�, 2007 (Part 3, th�s �ssue) from the northern Tasman Sea, S. crassispinus Last, Edmunds and Yearsley, 2007 (Part 2, th�s �ssue) and S. megalops (Macleay, 1881) from off Austral�a.

METHODS

Methods follow those outl�ned �n Part 1 of th�s �ssue (Last et al., 2007c). The only mater�al, the holotype and two paratypes, of the new spec�es were measured �n full (Table 1). In the descr�pt�on, morphometr�c and mer�st�c values for the holotype are given first followed in parentheses by the ranges of the paratypes. Mer�st�cs were

taken from rad�ographs of all types of the new spec�es. Comparat�ve data for closely related Austral�an and New Caledon�an members of the ‘megalops-cubens�s group’ were obta�ned from data presented �n Parts 2 and 3 of th�s volume (Last et al., 2007a, 2007b).

Type spec�mens are depos�ted �n the �chthyolog�cal collect�on of the Museum of New Zealand, Well�ngton (NMNZ); their registration numbers are prefixed with th�s acronym. Comparat�ve data for 5 spec�mens of Squalus brevirostris Tanaka, 1917, a Japanese member of th�s spec�es group, are also presented. These spec�mens are depos�ted �n the Kagosh�ma Un�vers�ty Museum, Kagosh�ma, Japan (KAUM), and the Austral�an Nat�onal F�sh Collect�on, Hobart (CSIRO). Spec�men reg�strat�on numbers are prefixed by these acronyms.

Squalus raoulensis sp. nov.


Holotype. NMNZ P 41678, adult male 651 mm TL, off Raoul Island, Kermadec Islands, New Zealand, 29°14′ S, 177°53′ W, 320 m, 8 November 2004.Paratypes. 2 spec�mens. NMNZ P 42572, adult male 681 mm TL, collected w�th holotype; NMNZ P 34436, female 729 mm TL, Kermadec R�dge north of Raoul Island.

DIAGNOSIS.— A moderate-s�zed spec�es of Squalus of the ‘megalops-cubens�s group’ w�th the follow�ng comb�nat�on of characters: mouth w�dth 2.4–2.7 t�mes

32

Figure 1. Lateral v�ew of Squalus raoulensis sp. nov. holotype (NMNZ P 41678, adult male 651 mm TL).

Figure 2. Ventral v�ew of the head of Squalus raoulensis sp. nov. holotype (NMNZ P 41678, adult male 651 mm TL).

hor�zontal prenar�al length; d�rect pre-second dorsal length 3.6–4.0 times pectoral-fin anterior margin, 2.7–2.9 t�mes dorsal caudal marg�n; preoral length 3.0–3.2 t�mes hor�zontal prenar�al length, 2.3–2.5 t�mes �nternar�al width; anterior nasal flap bifurcate; first dorsal fin upright and rounded apically; dorsal-fin spines not robust, width of exposed bases 0.6–0.7% TL; first dorsal-fin height 1.2–1.4 times its inner margin length; second dorsal-fin he�ght 0.85–0.93 t�mes �ts �nner marg�n length; pectoral-fin anterior margin 1.7–1.8 times its inner margin length; preventral caudal marg�n 1.7–2.3 t�mes �nner marg�n of pelvic fin; caudal fin pale with strongly demarcated, whitish posterior margin, no caudal bar; flank denticles un�cusp�d; 41–43 monospondylous centra, 84–85 precaudal centra, 112–113 total centra; adult max�mum s�ze at least 73 cm TL.

DESCRIPTION.— Body fus�form, slender, nape not prom�nently humped, only marg�nally deeper at belly (un�formly deepest across abdomen), max�mum depth 0.97 (0.93–1.00 �n paratypes) t�mes w�dth; trunk depth 1.04 (1.02–1.08 �n paratypes) t�mes abdomen depth; head short 22.5 (22.3–22.9)% TL; caudal peduncle slender, 26 (23.2–26.4)% TL. Head not espec�ally broad, w�dth 1.24 (1.17–1.22) t�mes trunk w�dth, 1.37 (1.35–1.53) t�mes abdomen w�dth, sl�ghtly depressed forward of spiracles, becoming subtriangular towards pectoral-fin or�g�n, length 2.1 (2.14–2.17) �n pre-vent length; he�ght 0.75 (0.73–0.74) t�mes w�dth. Snout short, narrowly tr�angular �n lateral v�ew, apex po�nted, lateral prenar�al marg�n somewhat angular; bluntly po�nted �n dorsal v�ew, hor�zontal length 1.3 (1.19–1.25) t�mes eye length, 0.7 (0.65–0.67) t�mes �nterorb�tal length; hor�zontal


prenar�al length 3.15 (3.03–3.11) t�mes �n preoral length. Eye broadly oval, s�ze moderate, length 4.95 (4.74–4.99) �n head, 2.07 (1.97–2.74) t�mes he�ght; strongly notched poster�orly, weakly connected to anteroventral marg�n of sp�racle. Sp�racle small, strongly crescent�c; lobe-l�ke fold on poster�or marg�n; a narrow subvert�cal fold present at anter�or marg�n �n holotype, fold covered �n dent�cles (less obv�ous �n female paratype NMNZ P 34436), greatest d�ameter 3.9 (3.51–4.28) �n eye length. G�ll open�ngs sl�ghtly obl�que, d�rected sl�ghtly anterodorsally from bottom to top (somet�mes upr�ght), first four subequal in size, fifth longest, height of fifth slit 1.8 (1.76–1.96)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.23 (1.12–1.32) �n preoral length; upper lab�al furrow more than tw�ce length of lower furrow, prom�nent postoral groove, sl�ghtly longer than upper lab�al furrow, extend�ng posterolaterally from angle of jaws; two ser�es of funct�onal teeth �n upper and lower jaws. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Nostr�ls small, sl�ghtly oblique; anterior nasal flap moderately bifurcate, upper lobe largest, broadest; poster�or lobe short, d�g�t�form; �nternar�al space 2.26 (2.29–2.45) �n preoral length, 2.52 (2.28–2.38) t�mes nostr�l w�dth. Dermal dent�cles on flank very small, unicuspid, not imbricated; crown quadrangular, w�dth subequal to or sl�ghtly narrower than length, w�th pronounced med�an r�dge; med�an r�dge commenc�ng well anter�or of dent�cle base w�th a mes�al furrow develop�ng anter�orly and converg�ng towards poster�or t�p of crown; cusp pungent, elongate; lateral extens�ons each w�th a weak r�dge, cusps absent. F�rst dorsal fin somewhat upright, broadly rounded apically; anter�or marg�n weakly convex; poster�or marg�n weakly concave, greatest curvature at or just below m�d-length; free rear t�p very th�ck basally, short, �nner marg�n of fin almost straight; insertion of base extremely well forward of pelvic-fin origin, just posterior to free rear

Figure 3. Cusps of the flank denticles of Squalus raoulensis sp. nov. paratype (NMNZ P 42572, adult male 681 mm TL). F�eld of v�ew w�dth 0.4 mm.

tip of pectoral fin; fin-spine origin well behind pectoral-fin insertion; spine base moderately robust, exposed anter�orly just below junct�on of sp�ne and soft port�on of fin; soft portion of fin connected distally at about m�d-po�nt of total sp�ne length; sp�ne taper�ng un�formly d�stally, anter�or marg�n almost stra�ght, subequal �n length to second dorsal-fin spine, tip well below apex of soft portion of fin in undamaged holotype; pre-first dorsal length 3.39 (3.36–3.39) times in TL; first dorsal-fin length 1.75 (1.97–2.00) times its height, 1.02 (1.13) times second dorsal-fin length; first dorsal-fin height 1.62 (1.57–1.59) times second dorsal-fin height; exposed first dorsal-spine length 0.51 (0.36) times height of fin. Second dorsal fin of moderate size, not strongly raked; anter�or marg�n weakly convex, apex narrowly rounded; poster�or marg�n deeply concave, max�mum concav�ty sl�ghtly near m�d-po�nt of marg�n, upper port�on almost upr�ght (d�rected sl�ghtly dorsoposter�orly from bottom to top �n paratype NMNZ P 34436); free rear t�p th�n basally, elongate, �nner marg�n length 1.07 (1.15–1.17) times fin height; second dorsal-fin length 2.79 (2.73–2.8) times its height; spine length 0.99 (0.98) in height of fin; fin-spine origin well behind free rear tip of pelvic fin; �nterdorsal space 0.97 (0.90–0.91) �n length from snout t�p to pectoral-fin origin, 1.27 (1.25–1.26) in pre-first dorsal length; second sp�ne moderately robust, not extremely broad-based, subequal to width of exposed base of first dorsal-fin spine; spine slender, sharply pointed distally, taper�ng gradually just above po�nt of exposure, t�p just below level of apex of soft portion of fin; no obvious

Figure 4. Lateral view of the dorsal fins of Squalus raoulensis sp. nov. holotype (NMNZ P 41678, adult male 651 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B

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Table 1. Proport�onal d�mens�ons as percentages of total length for the holotype (NMNZ P 41678) and ranges for the two paratypes of Squalus raoulensis sp. nov.

S. raoulensis sp. nov. Holotype Paratypes

M�n. Max.TL – Total length 651 681 729PCL – Precaudal length 78.5 78.2 78.6PD2 – Pre-second dorsal length 59.4 60.1 61.3PD1 – Pre-first dorsal length 29.5 29.5 29.8SVL – Pre-vent length 47.3 47.9 49.7PP2 – Prepelv�c length 45.2 46.5 48.8PP1 – Prepectoral length 22.3 21.4 21.4HDL – Head length 22.5 22.3 22.9PG1 – Prebranch�al length 18.8 18.2 18.7PSP – Presp�racular length 13.0 13.3 13.7POB – Preorb�tal length 7.2 7.1 7.3PRN – Prenar�al length 4.4 4.2 4.4POR – Preoral length 9.6 8.9 9.6MOW – Mouth w�dth 7.8 7.3 7.9ULA – Lab�al furrow length 2.4 2.0 2.3INW – Internar�al space 4.2 3.9 3.9INO – Interorb�tal space 8.5 8.6 8.7EYL – Eye length 4.5 4.5 4.8EYH – Eye he�ght 2.2 1.8 2.3SPL – Sp�racle length 1.2 1.1 1.3GS1 – F�rst g�ll-sl�t he�ght 1.8 1.8 1.8GS5 – F�fth g�ll-sl�t he�ght 1.8 1.8 2.0IDS – Interdorsal space 23.1 23.6 23.7DCS – Dorsal-caudal space 10.8 9.6 10.0PPS – Pectoral-pelv�c space 21.3 22.0 22.0PCA – Pelv�c-caudal space 26.0 23.2 26.4D1L – F�rst dorsal length 13.9 14.4 14.8D1A – F�rst dorsal anter�or marg�n 12.2 11.8 12.1D1B – F�rst dorsal base length 8.2 8.3 8.4D1H – F�rst dorsal he�ght 8.0 7.2 7.5D1I – F�rst dorsal �nner marg�n 5.9 6.3 6.3D1P – F�rst dorsal poster�or marg�n 8.8 9.1 9.3D1ES – F�rst dorsal sp�ne length 4.0 2.6 2.7D1BS – F�rst dorsal sp�ne base w�dth 0.7 0.6 0.7D2L – Second dorsal length 13.7 12.7 13.1D2A – Second dorsal anter�or marg�n 11.6 10.1 10.5D2B – Second dorsal base length 8.4 7.2 7.7D2H – Second dorsal he�ght 4.9 4.5 4.8D2I – Second dorsal �nner marg�n 5.3 5.2 5.6D2P – Second dorsal poster�or marg�n 5.5 5.4 6.2D2ES – Second dorsal sp�ne length 4.9 – 4.7D2BS – Second dorsal sp�ne base w�dth 0.8 0.7 0.7


Table 1. cont’d

S. raoulensis sp. nov. Holotype Paratypes

M�n. Max.P1A – Pectoral anter�or marg�n 15.3 15.0 16.9P1I – Pectoral �nner marg�n 9.1 8.8 9.1P1B – Pectoral base length 5.1 5.1 5.4P1P – Pectoral poster�or marg�n 11.8 12.4 13.9P2L – Pelv�c length 11.5 10.1 10.6P2H – Pelv�c he�ght 5.0 5.0 5.3P2I – Pelv�c �nner marg�n 6.3 5.0 6.4CDM – Dorsal caudal marg�n 22.0 20.9 21.4CPV – Preventral caudal marg�n 11.5 11.3 11.4CPU – Upper postventral caudal marg�n 15.9 15.8 16.0CPL – Lower postventral caudal marg�n 5.4 4.9 5.2CFW – Caudal fork w�dth 7.1 7.1 7.7CFL – Caudal fork length 9.8 9.7 10.0HANW – Head w�dth at nostr�ls 6.9 6.6 6.6HAMW – Head w�dth at mouth 11.4 11.1 11.9HDW – Head w�dth 12.7 13.3 13.5TRW – Trunk w�dth 10.3 10.9 11.5ABW – Abdomen w�dth 9.3 8.7 10.0TAW – Ta�l w�dth 7.0 6.2 7.0CPW – Caudal peduncle w�dth 3.0 3.2 3.3HDH – Head he�ght 9.6 9.8 9.8TRH – Trunk he�ght 10.0 10.7 10.8ABH – Abdomen he�ght 9.6 10.1 10.5TAH – Ta�l he�ght 7.1 6.5 7.2CPH – Caudal peduncle he�ght 2.5 2.4 2.7CLO – Clasper outer length 3.5 – 3.1CLI – Clasper �nner length 9.7 – 9.8CLB – Clasper base w�dth 1.6 – 1.5

�nterdorsal r�dge or groove (weak �n paratype NMNZ P 34436). Pectoral fin moderately elongate, anterior margin moderately convex; �nner marg�n moderately convex, length 9.1 (8.8–9.1)% TL; apex narrowly rounded to po�nted, lobe-l�ke but not falcate; poster�or marg�n moderately to strongly concave, free rear t�p usually angular; base 3.0 (2.95–3.11) �n length of anter�or marg�n. Pelvic fins small, anterior and posterior margins almost stra�ght (poster�or marg�n sl�ghtly convex �n female paratype NMNZ P 34436), apex broadly rounded, free rear t�p angular (more obtuse �n female paratype). Caudal peduncle long, tapering evenly to caudal fin; subcircular �n cross-sect�on anter�orly, sl�ghtly depressed poster�orly, ventral groove very well developed w�th weak med�an r�dge; lateral keels well developed, or�g�nat�ng under insertion of second dorsal fin, terminating much less than an eye diameter behind caudal-fin insertion; pelvic–

caudal space 0.82 (0.83–0.95) �n pectoral–pelv�c space, 0.86 (0.81–0.92) �n prepectoral length; dorsal–caudal space 2.15 (2.36–2.46) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal p�t weak. Caudal fin of moderate size, upper lobe not especially broad, upper postventral marg�n moderately convex; lower lobe well developed, narrowly rounded (angular �n female paratype); dorsal caudal marg�n 1.03 (1.04–1.10) �n head length; length of lower caudal lobe 1.92 (1.86–1.88) �n upper lobe length. Vertebral centra 112 (112–113 �n paratypes), monospondylous 41 (41–43), precaudal 84 (85) and caudal 28 (27–28). Teeth �n upper jaw 13+13 (13+13)=26, lower jaw 12+12 (11–12 + 12)=24 (23–24).

COLOUR.— When fresh: un�form redd�sh brown�sh dorsally; much paler, wh�te on ventral surface; l�ght and

36

dark areas well d�fferent�ated; sharply demarcated border extend�ng from snout t�p, below eye, through m�ddle of gill slits to pectoral fin origin; light and dark areas merging gradually on abdomen and tail; fins dark reddish brown. Dorsal-fin spines dusky, darkest anteriorly; base of soft portion slightly paler than distal portion of fin; first dorsal fin with very narrow black apical margin, extending to about d�stal th�rd of poster�or marg�n, lower two th�rds of poster�or marg�n and free rear t�p w�th narrow wh�te edge; second-dorsal fin similar colour to first dorsal fin, dark edge confined to fin tip, entire posterior margin with wh�te edge, ventral marg�n of free rear t�p brown�sh. Caudal fin mostly reddish brown; sharply defined white border extending along entire posterior margin of fin, its width subequal to half length of fifth gill slit. Pectoral and pelvic fin upper surfaces brownish, ventral surfaces pale, whitish; prominent, sharply defined white margins along posterior and inner margins of pectoral fin, and posterior margin of pelvic fin; claspers almost uniformly wh�te. In preservat�ve: colorat�on s�m�lar to above except redd�sh brown areas becom�ng grey�sh brown; pale and darker areas on dorsal fins remaining strongly contrasted; dark areas on upper surfaces of pectoral fin evident from ventral surface, most evident on lateral distal half of fin; mouth white; naked axils of fins white or dusky.

SIZE.— Females atta�n at least 729 mm TL; males probably smaller, matur�ty s�ze unknown, male types adult at 651 and 681 mm TL.

DISTRIBUTION.— Known only from the upper �nsular slope west of Nap�er Island (off Raoul Island), and north of Raoul Island on the Kermadec R�dge. Not collected from the Kermadec R�dge south of Raoul Island.

ETYMOLOGY.— The ep�thet raoulensis �s der�ved �n reference to the type local�ty of Raoul Island.

VERNACULAR.— Kermadec Spiny Dogfish.

REMARKS.— Squalus raoulensis and the larger and more w�dely-d�str�buted New Zealand spurdog S. griffini, are sympatr�c at Raoul Island �n the northern sector of the Kermadec group. Squalus raoulensis can be read�ly d�st�ngu�shed from S. griffini by �ts un�cusp�d dent�cles (rather than tr�cusp�d �n S. griffini), shorter snout (preorb�tal length 7.1–7.3 vs. 7.4–8.8% TL), lower vertebral count (112–113 vs. generally 116–117, max. 121), and smaller s�ze at matur�ty (males mature by 651 mm TL vs. 690– 760 mm TL). Of members of the ‘megalops-cubens�s group’, �ts closest reg�onal relat�ve �s S. bucephalus from nearby New Caledon�a. Squalus raoulensis ma�nly d�ffers from th�s spec�es �n hav�ng a narrower head (max�mum w�dth 12.7–13.5 vs. 13.7–14.2% TL, w�dth at anter�or nostr�ls 6.6–6.9 vs. 7.7–9.3% TL �n S. bucephalus), more slender body (trunk w�dth 10.3–11.5 vs. 11.8–12.9% TL, trunk he�ght 10.0–10.8 vs. 11.5–13.1% TL), and fewer vertebrae (41–43 vs. 45 monospondylous centra; 84–85 vs. 86–89 precaudal centra; and 112–113 vs.

113–119 total centra). It also has a shorter d�rect outer prenar�al length (4.2–4.4 vs. 4.6–5.2% TL) and upper postventral caudal marg�n (15.8–16.0 vs. 17.1–18.3% TL), a relatively smaller first dorsal fin (first dorsal-fin base 8.2–8.4 vs. 9.1–10.1% TL, he�ght 7.2–8.0 vs. 8.1–8.5% TL), a narrower exposed first dorsal-fin spine base (width of exposed base of first dorsal-fin spine 0.6–0.7 vs. 0.8% TL), a longer pre-first dorsal length (29.5–29.8 vs. 27.7–28.8% TL) and �nner nostr�l-upper lab�al furrow space (5.2–5.4 vs. 4.5–5.0% TL), and a relat�vely larger second dorsal fin (second dorsal-fin length 12.7–13.7 vs. 11.0–12.7% TL, second dorsal-fin inner margin 5.2–5.6 vs. 4.0–5.0% TL).

Squalus raoulensis d�ffers �n morphology and mer�st�cs from the Austral�an members of the ‘megalops-cubens�s group’, S. crassispinus and S. megalops. Its dorsal-fin sp�ne bases are more slender than S. crassispinus (0.6–0.7 vs. 1.2–1.5% TL), �ts mouth w�dth 2.4–2.7 t�mes (rather than 2.1–2.4 t�mes �n S. crassispinus, n=7) the hor�zontal prenar�al length, preoral length 3.0–3.2 t�mes the horizontal prenarial length (rather than 2.7–3.0), the first dorsal fin more upright (first dorsal-fin height 1.2–1.4 vs. 1.3–1.7 times its inner margin length), second dorsal-fin he�ght 0.9 t�mes �ts �nner marg�n length (rather than 1.1–1.3), pectoral-fin anterior margin 1.7–1.8 times its inner marg�n length (rather than 1.9–2.2), preventral caudal margin 1.8–2.3 times the inner pelvic-fin margin (rather than 2.4–4.6), and the white caudal fin posterior margin �s strongly demarcated (rather than poorly demarcated). Compared to S. megalops from southeastern Austral�a (n=6), �t has a narrower �nterdorsal space (23.1–23.7 vs. 24.0–25.3% TL), smaller gills (first gill-slit height 1.8 vs. 2.0–2.4% TL), and larger prenar�al spaces (4.2–4.4 vs. 3.7–4.1% TL) and fins (pectoral-fin inner margin 15.0–16.9 vs. 13.6–14.9% TL; lower postventral caudal margin 4.9–5.2 vs. 4.0–4.6% TL; second dorsal-fin length 12.7–13.7 vs. 11.0–12.7% TL). The two spec�es can be unamb�guously d�st�ngu�shed by the rat�o, preoral length/�nternar�al w�dth (2.3–2.5 vs. 1.9–2.2 �n populat�ons of S. megalops from southeastern Austral�a, 2.0–2.1 from Queensland, and 2.0–2.1 from Western Austral�a). Squalus raoulensis has more vertebrae (112–113, mean 112.3) than both S. crassispinus (107–111, mean 109.4, n=11) and S. megalops (102–110, mean 105.3, n=41).

Squalus raoulensis superficially resembles S. brevirostris Tanaka, 1917 from southern Japan (F�gs 5 and 6). It can be d�st�ngu�shed from that spec�es by the proport�ons of the head and gills, size and shape of the dorsal fins, colorat�on and dent�cle morphology. Squalus raoulensis has a longer head (head length 22.3–22.9 vs. 20.3–21.2% TL �n S. brevirostris) and generally longer snout (pre-orbital length 7.1–7.3 vs. 6.6–7.1% TL), smaller gills (fifth gill height 1.8–2.0 vs. 2.2–2.8% TL), larger first (height 7.2–8.0 vs. 6.1–6.5% TL) and second dorsal fins (height 4.5–4.9 vs. 3.3–3.8% TL), second dorsal fin larger relative to the first (first dorsal height/second dorsal height about 1.6 vs. 1.7–1.9) and not strongly raked. In S. brevirostris


Figure 5. Lateral v�ew of Squalus brevirostris (KAUM I 184, female 540 mm TL).

Figure 6. Ventral v�ew of the head of Squalus brevirostris (KAUM I 184, female 540 mm TL).

the posterior margin of the second dorsal fin is deeply concave, and the apex of the second dorsal sp�ne �s located over or sl�ghtly beyond the notch �n the poster�or marg�n (cf. over or slightly beyond fin insertion in S. raoulensis). In S. raoulensis the dark dorsal colorat�on extends well down the s�de of the head, reach�ng the m�ddle of the gills, and onto the pelvic-fin base and upper surface of the pelvic fin. In S. brevirostris the dorsal colorat�on �s less extens�ve, only extend�ng to the top of the g�lls, and the pelvic-fin base and upper pelvic fin surfaces are pale, s�m�lar �n colour to the ventral abdomen (Pl. 364 Tanaka 1917). Adult S. raoulensis have un�cusp�d lateral dermal dent�cles wh�ch are arrow-shaped �n S. brevirostris. Tanaka (1917) descr�bed the dent�cles of the 430 mm male holotype as hav�ng “a sharp med�an keel and a short blunt one on e�ther s�de at a r�ght angle to the former at the base, forming more or less a cross figure” (�.e. un�cusp�d). As the holotype �s cons�derably smaller than mater�al exam�ned, shape d�fferences observed are l�kely to be ontogenet�c (see Garr�ck 1960).

Another large Atlant�c member of the ‘megalops-cubens�s group’, Squalus cubensis, has a shorter predorsal length (27.6–28.2 vs. 29.5–29.8% TL �n S. raoulensis), longer

snout (prenar�al length 4.6–4.8 vs. 4.2–4.4% TL), longer caudal fin (dorsal caudal margin 22.1–23.0 vs. 20.9–22.0% TL), and lower second dorsal fin (height 3.8 vs. 4.5–4.9% TL) (see Last et al., 2007a, Part 2 of th�s �ssue).

Other material. Squalus brevirostris: KAUM 184, female 538 mm TL, KAUM 186, female 552 mm TL, KAUM 187, female 590 mm TL, KAUM 190, female 578 mm TL, KAUM 191, female 537 mm TL, CSIRO H 6483–01, adult male 451 mm TL, CSIRO H 6483–02, adult male 452 mm TL, CSIRO H 6483–03, female 542 mm TL, CSIRO H 6483–04, female 514 mm TL, East Ch�na Sea off Kasasa, M�nam�satsuma, Kagosh�ma, Japan, 31°29′ N, 130°02′ E, 145–150 m.

ACKNOWLEDGEMENTS

Th�s project was part�ally funded by the New Zealand Department of Conservat�on, New Zealand B�od�vers�ty Strategy, and B�osystemat�cs of New Zealand EEZ F�shes (FRST Contract MNZ0302 to Te Papa). In part�cular, we thank Cl�ve Roberts (NMNZ) for support�ng th�s project. The holotype and male paratype were collected from the Kermadec Islands Mar�ne Reserve under a permit to undertake specified scientific research issued by the Auckland Conservancy, New Zealand Department of Conservat�on. Malcolm Franc�s, Andrew Stewart, L�ndsay Chadderton and the crew of the FV Southern Salvor (Robert, Ph�l, John, Denn�s and Sacha), Stoney Creek Sh�pp�ng Co., ass�sted w�th the collect�on of spec�mens. Andrew Stewart (NMNZ) also photographed and t�ssue sampled the holotype and male paratype when they were landed, Carl Struthers (NMNZ) ass�sted w�th the curat�on, exam�nat�on and photography of the spec�mens, Adr�an Turner and Cather�ne Hobb�s (Un�vers�ty of Auckland) prepared and prov�ded SEMs, and Thomas Schultz (NMNZ) prepared rad�ographs. Toru M�yash�ta collected all of the Squalus brevirostris spec�mens exam�ned, and donated some of these to the Austral�an Nat�onal F�sh Collect�on. H�royuk� Motomura, curator of the Kagosh�ma Un�vers�ty Museum collect�on,

38

loaned us the KAUM mater�al. Thanks also go to Lou�se Conboy for ed�t�ng and prepar�ng �mages, Dan Gledh�ll for prepar�ng the l�terature summary, and W�ll�am Wh�te, Gordon Yearsley and John Pogonosk� for the�r ed�tor�al comments.

REFERENCES

Duffy, C.A.J. and Last, P.R. (2007) Part 9 — Redescr�pt�on of the Spiny Dogfish Squalus griffini Ph�ll�pps, 1931 from New Zealand, p. 91–100. In: Descr�pt�ons of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Garr�ck, J.A.F. (1960) Stud�es on New Zealand Elasmobranch��. Part XII. The spec�es of Squalus from New Zealand and Austral�a; and a general account and key to the New Zealand Squalo�dea. Transactions of the Royal Society of New Zealand, 88, 519–557.

Last, P.R., Edmunds, M. and Yearsley, G.K. (2007a) Part 2 — Squalus crassispinus sp. nov., a new spurdog of the ‘megalops-cubens�s group’ from the eastern Ind�an Ocean, p. 11–22. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Last, P.R., Séret, B. and Pogonosk�, J.J. (2007b) Part 3 — Squalus bucephalus, a new short-snout spurdog from New Caledon�a, p. 23–29. In: Descr�pt�ons of new

dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Last, P.R., Wh�te, W.T., Pogonosk�, J.J., Gledh�ll, D.C., Ward, B. and Yearsley, G.K. (2007c) Part 1 — Appl�cat�on of a rap�d taxonom�c approach to the genus Squalus, p. 1–10. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.



Paul�n, C., Stewart, A., Roberts, C. and McM�llan, P. (1989) New Zealand fish: a complete guide. GP Books: Well�ngton, 279 pp.

Phillipps, W.J. (1931) New species of piked dogfish. New Zealand Journal of Science and Technology, 12(6), 360–361.

Tanaka, S. (1917) Figures and descriptions of the fishes of Japan. Daiichi Shoin, Tokyo, 26, 455–474.


Part 5 — New species of Squalus of the ‘highfin megalops group’ from the Australasian region

Peter R. Last, W�ll�am T. Wh�te and John D. Stevens


ABSTRACT.— Three new spec�es of spurdogs, Squalus albifrons, S. altipinnis, and S. notocaudatus, are descr�bed based on mater�al from the outer cont�nental shelf and cont�nental slope of Austral�a. These spec�es belong to a subgroup of Squalus, known as the ‘highfin megalops group’, which have a short snout and white caudal-fin margin reminiscent of S. megalops. However, unl�ke S. megalops, they have a strong-spined, upright first dorsal fin and tricupsid denticles. Squalus altipinnis �s known from two spec�mens collected �n about 300 m depth near the Rowley Shoals, off northwestern Austral�a. Squalus albifrons and S. notocaudatus appear to be endem�c to the cont�nental shelf and slope off eastern Austral�a. Squalus albifrons occurs w�dely from central New South Wales to central Queensland �n 135–450 m depth. Squalus notocaudatus �s known from the upper cont�nental slope off central Queensland �n 225–450 m depth. The species can be distinguished from each other based on meristics, morphometrics, denticle shape, and fin colorat�on.

Key words. Squalo�dea – Squal�dae – Squalus albifrons – Squalus altipinnis – Squalus notocaudatus – new species – Indo–West Pacific


INTRODUCTION

Last and Stevens (1994) identified 9 Australian Squalus spec�es of wh�ch 5 rema�n formally undescr�bed. Three of these spec�es belong to a subgroup prov�s�onally called the ‘highfin megalops group’. Australian members of th�s group are character�sed by a short snout (preorb�tal length 6.4–7.0% TL), a tall and upright first dorsal fin (height 7.7–9.4% TL), robust dorsal-fin spines (exposed base w�dth 0.7–1.1% TL), tr�cusp�d dent�cles, and a white posterior margin on the caudal fin. Members of the ‘megalops-cubens�s group’ also have a short snout and white posterior margin on the caudal fin, but have d�st�nct�ve un�cusp�d dent�cles and more strongly raked dorsal fins. The new members of the ‘highfin megalops group’, informally identified as Squalus sp. A, Squalus sp. B and Squalus sp. C (sensu Last and Stevens, 1994), are descr�bed based on mater�al from Austral�a and eastern Indones�a. Other undescr�bed members of th�s subgroup may exist in the Indo-Pacific region.

METHODS

Methods follow those outl�ned �n Part 1 of th�s �ssue (Last et al., 2007). For Squalus albifrons sp. nov., both morphometr�cs and mer�st�cs were taken from the holotype (CSIRO H 4627–01) and the follow�ng 5 paratypes: CSIRO H 644–04, CSIRO H 2487–01, CSIRO H 4704–02, CSIRO H 4705–01 and QM I 38077.

In add�t�on, mer�st�cs were taken from the follow�ng 6 paratypes: CSIRO H 449, CSIRO H 2487–02, CSIRO H 2487–03, CSIRO H 2691–01, CSIRO H 3589–01 and QM I 19327. For Squalus altipinnis sp. nov., morphometr�cs and mer�st�cs were taken from the holotype (CSIRO CA 4111) and paratype (CSIRO CA 3297). For Squalus notocaudatus sp. nov., both morphometr�cs and mer�st�cs were taken from the holotype (CSIRO H 1368–02) and 3 paratypes (CSIRO H 1321–01, CSIRO H 1322–01 and CSIRO H 1323–01) (Table 1). In the spec�es descr�pt�ons, morphometr�c and meristic values for the holotypes are given first, followed �n parentheses by ranges for paratypes. Type spec�mens are depos�ted �n the Austral�an Nat�onal F�sh Collect�on, Hobart (CSIRO) and the �chthyolog�cal collect�on of the Queensland Museum, Br�sbane (QM); the�r reg�strat�on numbers are prefixed with these acronyms.

Squalus albifrons sp. nov.


Squalus sp. B: Last and Stevens, 1994, Sharks and Rays of Australia, pp 49, 93, figs 20, 8.32, pl. 6; Compagno, Dando and Fowler, 2005, Sharks of the World, p 79, figs, pl. 2. Holotype. CSIRO H 4627–01, adult male 618 mm TL, east of Broken Bay, New South Wales, 33°28′ S, 152°05′ E, 386 m, 18 Jun 1996.

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Paratypes. 11 spec�mens. CSIRO H 644–04, adult male 643 mm TL, Saumarez Reef, Queensland, 22°49′ S, 154°10′ E, 450 m; CSIRO H 449, adult male 650 mm TL, northeast of Townsv�lle Trough, Queensland, 17° 57′ S, 147°03′ E, 300 m; QM I 38077, adult male 614 mm TL, east of Cairns, Queensland, 17°11′ S, 146°41′ E, 260 m; QM I 19327, adult male 618 mm TL, off Noosa, Queensland, 26°20′ S, 153°53′ E, 275 m; CSIRO H 2691–01, adult male 695 mm TL, northeast of Byron Bay, New South Wales, 28°30′ S, 153°51′ E, 153 m; CSIRO H 2487–01, female 647 mm TL, CSIRO H 2487–02, female 658 mm TL, CSIRO H 2487–03, female 681 mm TL, southeast of Ball�na, New South Wales, 29°03′ S, 153°49′ E, 156 m; CSIRO H 3589–01, female 844 mm TL, east of Newcastle, New South Wales, 32°59′ S, 152°14′ E, 135 m; CSIRO H 4705–01, female 614 mm TL, east of Sydney, New South Wales, 33°28′ S, 152°00′ E, 230 m; CSIRO H 4704–02, female 596 mm TL, east of Sydney, New South Wales, 33°37′ S, 151° 55′ E, 331 m.Non-types. 3 spec�mens. CSIRO H 4704–01, female 656 mm TL, east of Sydney, New South Wales, 33°37′ S, 151°55′ E, 331 m; CSIRO H 3711–01, female 753 mm TL, northeast of Wollongong, New South Wales, 34°18′ S, 151°12′ E, 135 m; CSIRO H 4709–01, female 784 mm TL, east of Brush Island, New South Wales, 35°34′ S, 150°44′ E, 388 m.

DIAGNOSIS.— A large spec�es of Squalus of the ‘highfin megalops group’ with the following combination of characters: abdomen depth 10.4–12.6% TL; pre-vent length 46.9–48.1% TL, 2.0–2.2 t�mes dorsal caudal marg�n; pre-second dorsal length 3.7–4.3 t�mes pectoral-fin anterior margin, 2.5–2.8 times dorsal caudal margin; head w�dth 1.0–1.4 t�mes abdomen w�dth; preoral length 2.9–3.1 t�mes hor�zontal prenar�al length, 8.4–9.1% TL; head length 4.3–4.9 t�mes eye length; mouth w�dth 3.3–4.1 t�mes length of upper lab�al furrow; �nterorb�tal width 1.5–1.7 times horizontal preorbital length; fifth gill slit height 2.1–2.5% TL; anterior nasal flap strongly bifurcate; first dorsal fin upright, upper posterior margin d�rected posteroventrally, greatest concav�ty about midway between free rear tip and fin apex; posterior margin of second dorsal fin deeply concave; second dorsal-fin spine with a broad base; pectoral fin of adult not falcate, anter�or marg�n long, 14.2–15.8% TL; dorsal surface dark, sharply demarcated from paler ventral surface on side of head; dorsal fins dark with obvious darker tips; first dorsal-fin spine distinctly paler than base of soft portion of dorsal fin; caudal fin with broad white posterior margin, no caudal bar; flank denticles weakly tr�cusp�d; 44–46 monospondylous centra, 89–93 precaudal centra, 116–122 total centra; adult max�mum s�ze at least 84 cm TL.

DESCRIPTION.— Body fus�form, moderately elongate (some s�m�lar-s�zed paratypes more robust), nape prominently humped; deepest near first dorsal-fin spine, max�mum depth 0.96 (1.03–1.12 �n paratypes) t�mes w�dth;

trunk depth 1.05 (0.92–1.01) t�mes abdomen depth; head short 21.1 (20.4–21.6)% TL; caudal peduncle relat�vely robust, 25.6 (24.4–26.7)% TL. Head moderately robust, sl�ghtly w�der than trunk, w�dth 1.13 (1.08–1.27) t�mes trunk w�dth, 1.31 (1.03–1.38) t�mes abdomen w�dth, depressed forward of sp�racles, becom�ng subtr�angular towards pectoral-fin origin, length 2.24 (2.20–2.36) in pre-vent length; he�ght 0.82 (0.80–0.87) t�mes w�dth. Snout short, narrowly tr�angular �n lateral v�ew, apex bluntly po�nted, lateral prenar�al marg�n not angular (weakly angular �n some paratypes); broadly rounded �n dorsal v�ew (sl�ghtly more po�nted �n some paratypes), hor�zontal length 1.16 (1.16–1.35) t�mes eye length, 0.62 (0.58–0.67) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.97 (2.91–3.06) t�mes �n preoral length. Eye narrowly oval, s�ze moderate, length 4.42 (4.33–4.88) �n head length, 2.68 (2.01–2.79) t�mes eye he�ght; strongly notched poster�orly, notch extend�ng halfway to sp�racle as a well-developed furrow (fully connected to m�d-anter�or marg�n of sp�racle �n some paratypes). Sp�racle moderate, subtr�angular to crescent�c; broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.36 (2.73–4.30) in eye length. Gill openings almost upright, first four subequal in size, fifth longest, height of fifth slit 2.3 (2.1–2.5)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.05 (1.04–1.17) �n preoral length; upper lab�al furrows less than tw�ce length of lower furrows; prom�nent post-oral groove, subequal �n length to upper lab�al furrows, extend�ng posterolaterally from angle of jaws; two ser�es of funct�onal teeth �n upper jaw, two ser�es �n lower. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Nostr�ls small, almost transverse; anter�or nasal flap strongly bifurcate, upper lobe largest, broadest; posterior lobe narrow, finger-like; internarial space 2.08 (1.86–2.07) �n preoral length, 2.40 (2.28–2.82) t�mes nostr�l length. Dermal dent�cles (based on male paratypes CSIRO H 449 and CSIRO H 644–04) on flank very small, weakly to moderately tr�cusp�d w�th pronounced med�an r�dge; med�an r�dge commenc�ng well anter�or of rest of crown, w�th a mes�al furrow develop�ng anter�orly and converg�ng towards poster�or t�p of crown; lateral cusps weakly developed, blunt to rud�mentary poster�orly, r�dge extend�ng d�stally, almost to t�p of cusp, w�th a shallow mes�al furrow; crown of female paratype more strongly tr�cusp�d and more �mbr�cated than that of male paratype. First dorsal fin elevated, upright, narrowly rounded ap�cally; anter�or marg�n moderately convex; upper poster�or marg�n almost stra�ght, not vert�cal, d�rected well anterodorsally from bottom to top, weakly concave (somet�mes moderately concave) near free rear t�p; free rear t�p relat�vely th�ck basally, short (short to moderate �n paratypes); inner margin of fin almost straight; insertion of base extremely well forward of pelvic-fin origin, slightly posterior to free rear tip of pectoral fin; fin-spine origin above (sometimes just behind) pectoral-fin insertion; sp�ne base broad, exposed anter�orly near junct�on of spine and soft portion of fin; soft portion of fin connected


Figure 1. Lateral v�ew of: Squalus albifrons sp. nov. holotype (CSIRO H 4627–01, adult male 618 mm TL; �mage reversed).

Figure 2. Ventral v�ew of the head of Squalus albifrons sp. nov. holotype (CSIRO H 4627–01, adult male 618 mm TL).

near m�d-po�nt of total sp�ne length; sp�ne taper�ng rap�dly d�stally (less so �n some paratypes), anter�or marg�n sl�ghtly convex (somet�mes almost stra�ght); exposed port�on of sp�ne upr�ght to t�lted sl�ghtly, subequal �n length to exposed portion of second dorsal-fin spine; pre-first dorsal length 3.66 (3.49–3.76) times in TL; first dorsal-fin length 1.69 (1.67–1.77) times its height, 1.16 (1.15–1.21) times second dorsal-fin length; first dorsal-fin height 1.78 (1.64–1.86) times second dorsal-fin height; exposed first dorsal-spine length 0.55 (0.53–0.63) times height of fin. Second dorsal fin of moderate size, strongly raked; anter�or marg�n strongly convex, apex narrowly rounded; poster�or marg�n deeply concave, max�mum

concav�ty almost near m�d-po�nt of marg�n or sl�ghtly above, upper port�on d�rected dorsoposter�orly strongly from bottom to top; free rear t�p elongate, �nner marg�n length 1.01 (0.86–1.00) times fin height; second dorsal-fin length 2.59 (2.36–2.69) times its height; spine length 1.01 (0.85–1.09) in height of fin; fin-spine origin well behind free rear tip of pelvic fin, exposed at about level of junction with spine and soft portion of fin; second sp�ne moderately broad based, slender, sharply po�nted d�stally, taper�ng rap�dly (somet�mes evenly) from po�nt of exposure, sp�ne t�p extend�ng beh�nd level of insertion of fin (when undamaged), much more strongly raked than first spine; interdorsal space 0.87 (0.81–0.88)

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in pre-pectoral length, 1.12 (1.08–1.11) in pre-first dorsal length; moderate interdorsal groove. Pectoral fin moderate, anter�or marg�n weakly to moderately convex; �nner marg�n weakly convex, length 7.7 (6.4–7.6)% TL; apex moderately rounded, lobe-l�ke but not falcate; poster�or marg�n moderately concave, free rear t�p bluntly angular; base very short, 2.70 (2.56–2.92) �n length of anterior margin. Pelvic fins small, anterior and posterior marg�ns almost stra�ght, apex broadly rounded, free rear t�p acute. Caudal peduncle long, taper�ng sl�ghtly to caudal fin; subcircular in cross-section anteriorly, broadly sem�c�rcular poster�orly, ventral groove usually very well developed; lateral keels well developed, or�g�nat�ng under insertion of second dorsal fin (more posterior �n paratypes), term�nat�ng less than an eye d�ameter behind caudal-fin insertion; pelvic–caudal space 0.82 (0.73–0.91) �n pectoral–pelv�c space, 0.83 (0.79–0.86) �n prepectoral length; dorsal–caudal space 2.52 (2.29–2.59) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal pit rudimentary. Caudal fin relatively long, dorsal caudal marg�n 0.90 (0.86–1.01) �n head length; length of lower caudal lobe 1.95 (1.86–1.96) �n upper lobe length; upper poster�or lobe moderately convex; lower lobe apex somewhat angular (narrowly rounded �n some paratypes). Vertebral centra 119 (116–122 �n 10 paratypes), monospondylous 45 (44–46), precaudal 89 (90–93) and caudal 30 (26–31). Teeth �n upper jaw (�n paratypes CSIRO H 2487–02, CSIRO H 2487–03) 13–14+13–14=27, lower jaw 11–12+11=22–23.

COLOUR.— When fresh (based on holotype and CSIRO H 3589–01): dorsal half slate grey, wh�te ventrally; pale and dark tonal areas on head strongly demarcated before pectoral fin, their border extending from snout, under eye, through bottom half of g�ll sl�ts; demarcat�on on trunk less ev�dent, ta�l more un�formly grey ventrally. First dorsal fin dark greyish; holotype with a narrow black posterior margin extending from fin apex along the upper half of fin (not evident in paratype); free rear tip and base of fin dusky, not significantly paler than rest

Figure 3. Cusps of the flank denticles of Squalus albifrons sp. nov. paratype (CSIRO H 644–04, adult male 643 mm TL). F�eld of v�ew w�dth 1.2 mm.

of fin; second dorsal fin almost uniformly dusky, apex barely darker than rest of fin; first dorsal-fin spine pale; second dorsal-fin spine dusky with dark anterior margin; sk�n below exposed sp�ne bases d�st�nctly darker than fins. Caudal fin greyish, posterior margin of upper lobe wh�te; ventral lobe w�th a narrow, pale marg�n �n holotype (weaker �n paratype); base sl�ghtly darker than rest of fin. In preservative (based on holotype): body darker, more un�formly toned than �n fresh cond�t�on; d�st�nct pale and dark areas on head st�ll ev�dent; prom�nent white eyebrow above both eyes; pectoral and pelvic fins dark grey dorsally w�th narrow, pale poster�or marg�ns, fins similar ventrally apart from white basal portions; dorsal fins with persistent dark distal and upper posterior borders; free rear tips not paler than remaining fins; pale posterior margin of caudal fin persistent, bordered by a sl�ghtly darker submarg�nal bar.

SIZE.— Females reach�ng at least 844 mm TL (paratype); males probably smaller, largest male 695 mm TL (paratype), smallest adult male 614 mm TL (paratype). Females and males have been reported to atta�n 860 and 740 mm TL, respect�vely, off New South Wales (Graham, 2005).

DISTRIBUTION.— Known only from eastern Austral�a between the Townsv�lle Trough, Queensland (ca. 17° S), and off Ulladulla, southern New South Wales (ca. 35° S), �n 131–450 m depth.

Figure 4. Lateral view of the dorsal fins of: Squalus albifrons sp. nov. holotype (CSIRO H 4627–01, adult male 618 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B


ETYMOLOGY.— Der�ved from the Lat�n albi (wh�te) and frons (brow) �n allus�on to the wh�te upper ocular marg�n �n most spec�mens where dent�cles have been shed.

VERNACULAR.— Eastern Highfin Spurdog.

REMARKS.— Belongs to a group of spurdogs tentatively labeled as the ‘highfin megalops group’ that are d�st�ngu�shed by the comb�nat�on of a short snout, robust dorsal-fin spines, tricuspid denticles, a white posterior caudal-fin margin, and a tall, upright first dorsal fin. Of the Australian species, S. albifrons �s most s�m�lar morphometr�cally to another new spurdog descr�bed below �n th�s paper (�.e. S. sp. A: sensu Last and Stevens, 1994), but d�ffers �n the follow�ng characters: head length 4.3–4.9 (3.8–4.3 �n S. sp. A) t�mes eye length; �nterorb�tal space 1.48–1.73 (1.39–1.47) t�mes hor�zontal preorb�tal length; dorsal caudal marg�n 1.9–2.0 (2.0–2.1) t�mes pre-ventral caudal marg�n; pre-vent length 2.0–2.4 (1.9–2.0) times length of dorsal caudal margin; first dorsal-fin length 1.7–1.8 (1.5–1.7) times its height; fifth gill-slit length 2.1–2.5 (1.9–2.1)% TL; pelvic-fin inner margin length 5.0–6.0 (4.4–5.1)% TL; upper lab�al furrow 2.1–2.4 (1.8–2.1)% TL; and caudal peduncle w�dth 2.8–3.2 (2.1–2.6)% TL. Squalus albifrons, wh�ch appears to be more w�dely d�str�buted than the sympatr�c S. sp. A, also has a relat�vely darker, more contrasted, dorsal surface, and fewer vertebrae (89–93 vs. 94–97 precaudal centra). The posterior margin of the first dorsal fin of S. albifrons �s not strongly concave (rather than strongly concave) and the deepest part of the concav�ty �s near the m�ddle of the fin margin (rather than being situated more basally, closer to the free rear tip than the fin apex).

Squalus albifrons d�ffers from a th�rd new spurdog descr�bed �n th�s paper, (�.e. S. sp. C: sensu Last and Stevens, 1994), in having a larger caudal fin (dorsal caudal marg�n 21.3–23.6 vs. 19.6–19.9% TL �n S. sp. C; preventral caudal marg�n 11.2–12.5 vs. 10.8–11.4% TL; caudal fork length 9.4–10.5 vs. 8.4–8.5% TL, larger pectoral fin (anterior margin 14.2–15.8 vs. 13.9–14.1% TL; poster�or marg�n 10.8–11.7 vs. 10.2–10.4% TL) and shorter dorsal–caudal space (9.4–11.1 vs. 11.4–11.7% TL). Some other rat�os appear to be useful for d�st�ngu�sh�ng these spec�es: pre-second dorsal length 3.71–4.28 (vs. 4.32–4.36 �n S. sp. C) times pectoral-fin anter�or marg�n, 2.48–2.84 (vs. 3.07–3.08) t�mes dorsal caudal marg�n, 3.18–3.89 (vs. 4.16–4.22) t�mes upper postventral marg�n; mouth w�dth 3.29–4.08 (vs. 3.11–3.26) t�mes upper lab�al furrow; head length 2.52–2.73 (vs. 2.90–2.96) t�mes mouth w�dth; pre-vent length 2.00–2.24 (vs. 2.48) t�mes dorsal caudal marg�n; dorsal caudal marg�n 1.94–2.48 (vs. 1.70–1.72) t�mes dorsal–caudal space.

Squalus altipinnis sp. nov.


Squalus sp. C: Last and Stevens, 1994, Sharks and Rays of Australia, pp 49, 94, figs 21, 8.21, 8.33, pl. 6; Compagno, Dando and Fowler, 2005, Sharks of the World, p 79–80, figs, pl. 2.

Holotype. CSIRO CA 4111, adult male 586 mm TL, east of Rowley Shoals, Western Australia, 17°18′ S, 120° 09′ E, 305 m, 4 Feb 1984.Paratype. CSIRO CA 3297, adult male 589 mm TL, southwest of Rowley Shoals, Western Austral�a, 18° 10′ S, 118°20′ E, 298 m.

DIAGNOSIS.— A moderate-s�zed spec�es of Squalus of the ‘highfin megalops group’ with the following comb�nat�on of characters: abdomen depth 9.0–10.6% TL; pre-vent length 48.6–49.2% TL, about 2.5 t�mes dorsal caudal marg�n; pre-second dorsal length 4.3–4.4 times pectoral-fin anterior margin, about 3.1 times dorsal caudal marg�n; head w�dth about 1.5 t�mes abdomen w�dth; preoral length 2.9–3.2 t�mes hor�zontal prenar�al length, about 8.7% TL; head length 4.4–4.7 t�mes eye length; mouth w�dth 3.1–3.3 t�mes length of upper lab�al furrow; �nterorb�tal w�dth about 1.5 t�mes hor�zontal preorb�tal length; fifth gill slit height 2.2–2.5% TL; anterior nasal flap strongly bifurcate; first dorsal fin upright, upper poster�or marg�n d�rected posteroventrally, greatest concavity slightly closer to free rear tip than fin apex; posterior margin of second dorsal fin deeply concave; second dorsal-fin spine with a broad base; pectoral fin not falcate, anter�or marg�n short, 13.9–14.1% TL; dorsal surface sl�ghtly darker than ventral surface, but tones not sharply demarcated on side of head; dorsal fins pale with paler tips; first dorsal-fin spine darker than base of soft portion of dorsal fin; caudal fin with broad white posterior margin, no caudal bar; flank denticles weakly to moderately tr�cusp�d; 42–44 monospondylous centra, 88–92 precaudal centra, 114–120 total centra; adult max�mum s�ze at least 59 cm TL.

DESCRIPTION.— Body fus�form, slender, nape not prominently humped; barely deeper near first dorsal-fin sp�ne, max�mum depth 0.91 (0.90 �n paratype) t�mes w�dth, trunk depth 1.03 (0.89) t�mes abdomen depth; head short 22.0 (21.7)% TL; caudal peduncle slender, 27.1 (26.1)% TL. Head rather broad, much w�der than trunk, w�dth 1.23 (1.22) t�mes trunk w�dth, 1.53 (1.47) t�mes abdomen w�dth, moderately depressed forward of sp�racles, becom�ng weakly subtr�angular towards pectoral-fin origin, length 2.21 (2.27) in pre-vent length; he�ght 0.71 (0.77) t�mes w�dth. Snout short, tr�angular �n lateral v�ew, apex bluntly po�nted, lateral prenar�al marg�n not angular; broadly rounded �n dorsal v�ew, hor�zontal length 1.17 (1.23) t�mes eye length, 0.69 (0.67) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.92 (3.18) t�mes �n preoral length. Eye oval, s�ze moderate, length

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Figure 5. Lateral v�ew of: Squalus altipinnis sp. nov. holotype (CSIRO CA 4111, adult male 586 mm TL).

Figure 6. Ventral v�ew of the head of Squalus altipinnis sp. nov. holotype (CSIRO CA 4111, adult male 586 mm TL).

4.41 (4.72) �n head length, 2.76 (2.28) t�mes depth; strongly notched poster�orly, notch more pronounced anter�orly, extend�ng as a weak furrow to anteroventral marg�n of sp�racle. Sp�racle moderate, broadly crescent�c; broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.28 (2.93) in eye length. Gill openings almost upright, first four subequal in size, fifth longest, height of fifth slit 2.2 (2.5)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.17 (1.15) �n preoral length; upper lab�al furrows about tw�ce length of lower furrows (sl�ghtly less �n paratype); prom�nent postoral groove, subequal �n length to upper lab�al furrow, extend�ng posterolaterally from angle of jaws; two ser�es of funct�onal teeth �n upper jaw, two ser�es �n lower. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of

tooth broader than length of cusp. Nostr�ls small, almost transverse; anterior nasal flap strongly bifurcate, upper lobe largest, broadest; poster�or lobe narrow, somewhat thallate; �nternar�al space 2.13 (1.99) �n preoral length, 2.31 (2.48) t�mes nostr�l length. Dermal dent�cles (based on both types) on flank very small, weakly to moderately tr�cusp�date w�th pronounced med�an r�dge; med�an r�dge commenc�ng sl�ghtly anter�or of rest of crown, w�th a mes�al furrow develop�ng anter�orly and converg�ng gently towards poster�or t�p of crown; lateral cusps short or �nd�st�nct, blunt or broadly rounded poster�orly, r�dge extend�ng d�stally, almost to t�p of cusp, w�th low mesial furrow; strongly imbricated. First dorsal fin small, elevated, upr�ght, relat�vely broadly rounded ap�cally; anterior margin following profile of spine, strongly convex beyond sp�ne t�p; upper poster�or marg�n almost


Table 1. Proport�onal d�mens�ons as percentages of total length for the holotype (CSIRO H 4627–01) and ranges for the 5 paratypes of Squalus albifrons sp. nov. and the holotype (CSIRO CA 4111) and paratype (CSIRO CA 3297) of Squalus altipinnis sp. nov.

S. albifrons sp. nov. S. altipinnis sp. nov. Holotype Paratypes Holotype Paratype

M�n. Max.TL – Total length 618 596 647 586 589PCL – Precaudal length 76.4 75.9 78.1 79.9 80.0PD2 – Pre-second dorsal length 59.1 58.6 61.0 60.4 61.0PD1 – Pre-first dorsal length 27.3 26.6 28.6 27.6 27.0SVL – Pre-vent length 47.2 46.9 48.1 48.6 49.2PP2 – Prepelv�c length 45.0 45.0 46.3 46.1 46.5PP1 – Prepectoral length 21.3 20.2 21.8 22.2 21.7HDL – Head length 21.1 20.4 21.6 22.0 21.7PG1 – Prebranch�al length 17.9 17.2 18.1 18.4 18.0PSP – Presp�racular length 11.6 11.3 11.9 12.1 11.7POB – Preorb�tal length 6.9 6.7 7.0 6.8 6.8PRN – Prenar�al length 4.3 4.4 4.5 4.4 4.5POR – Preoral length 8.7 8.4 9.1 8.7 8.7INLF – Inner nostr�l-lab�al furrow space 4.5 4.4 4.8 4.7 4.7MOW – Mouth w�dth 8.3 7.5 8.6 7.4 7.5ULA – Lab�al furrow length 2.2 2.1 2.4 2.3 2.4INW – Internar�al space 4.2 4.3 4.6 4.1 4.3INO – Interorb�tal space 8.9 8.3 9.4 8.5 8.5EYL – Eye length 4.8 4.4 4.8 5.0 4.6EYH – Eye he�ght 1.8 1.7 2.2 1.8 2.0SPL – Sp�racle length 1.4 1.1 1.6 1.5 1.6GS1 – F�rst g�ll-sl�t he�ght 2.2 1.9 2.1 2.0 2.0GS5 – F�fth g�ll-sl�t he�ght 2.3 2.1 2.5 2.2 2.5IDS – Interdorsal space 24.4 24.4 26.1 23.8 24.8DCS – Dorsal-caudal space 9.7 9.4 11.1 11.4 11.7PPS – Pectoral-pelv�c space 21.1 19.5 22.6 21.8 22.1PCA – Pelv�c-caudal space 25.6 24.4 26.7 27.1 26.1D1L – F�rst dorsal length 14.6 13.7 14.9 14.7 14.9D1A – F�rst dorsal anter�or marg�n 12.7 10.9 12.5 12.6 12.0D1B – F�rst dorsal base length 8.9 8.5 9.3 9.6 9.6D1H – F�rst dorsal he�ght 8.6 7.7 8.9 7.8 7.9D1I – F�rst dorsal �nner marg�n 5.9 5.2 6.2 5.8 5.4D1P – F�rst dorsal poster�or marg�n 10.1 9.9 10.6 8.9 9.6D1ES – F�rst dorsal sp�ne length 4.8 4.4 5.4 4.9 5.3D1BS – F�rst dorsal sp�ne base w�dth 0.9 1.0 1.0 1.1 1.1D2L – Second dorsal length 12.5 11.4 12.9 12.9 12.6D2A – Second dorsal anter�or marg�n 11.1 10.0 11.3 11.5 10.9D2B – Second dorsal base length 7.7 7.3 8.3 8.5 7.9D2H – Second dorsal he�ght 4.8 4.3 5.1 4.5 4.8D2I – Second dorsal �nner marg�n 4.9 4.1 4.9 4.3 4.4D2P – Second dorsal poster�or marg�n 5.2 4.8 5.8 4.6 5.4D2ES – Second dorsal sp�ne length 4.9 3.8 5.2 4.7 4.8D2BS – Second dorsal sp�ne base w�dth 0.7 0.7 0.9 0.9 0.9

46

Table 1. cont’d

S. albifrons sp. nov. S. altipinnis sp. nov. Holotype Paratypes Holotype Paratype

M�n. Max.P1A – Pectoral anter�or marg�n 15.2 14.2 15.8 13.9 14.1P1I – Pectoral �nner marg�n 7.7 6.4 7.6 7.1 6.6P1B – Pectoral base length 5.6 5.4 6.0 5.2 5.1P1P – Pectoral poster�or marg�n 11.4 10.8 11.7 10.4 10.2P2L – Pelv�c length 10.6 9.2 9.8 10.4 10.0P2H – Pelv�c he�ght 5.1 4.3 4.7 4.7 4.9P2I – Pelv�c �nner marg�n 6.0 5.0 5.4 5.3 5.1CDM – Dorsal caudal marg�n 23.4 21.3 23.6 19.6 19.9CPV – Preventral caudal marg�n 12.0 11.2 12.5 10.8 11.4CPU – Upper postventral caudal marg�n 17.0 15.7 18.4 14.5 14.4CPL – Lower postventral caudal marg�n 4.4 3.9 4.9 4.0 4.3CFW – Caudal fork w�dth 6.5 6.8 7.3 6.4 6.8CFL – Caudal fork length 9.9 9.4 10.5 8.5 8.4HANW – Head w�dth at nostr�ls 6.7 6.6 7.1 6.8 6.7HAMW – Head w�dth at mouth 11.5 11.2 12.1 10.6 11.4HDW – Head w�dth 12.8 12.4 13.0 12.6 12.7TRW – Trunk w�dth 11.3 10.0 12.0 10.2 10.5ABW – Abdomen w�dth 9.8 9.4 12.5 8.3 8.6TAW – Ta�l w�dth 6.2 6.1 6.7 5.7 6.9CPW – Caudal peduncle w�dth 3.2 2.8 2.9 2.6 2.6HDH – Head he�ght 10.4 10.2 11.3 9.0 9.8TRH – Trunk he�ght 10.8 10.9 12.3 9.3 9.4ABH – Abdomen he�ght 10.4 11.2 12.6 9.0 10.6TAH – Ta�l he�ght 6.5 6.2 7.2 5.7 6.2CPH – Caudal peduncle he�ght 2.6 2.5 2.7 2.3 2.2CLO – Clasper outer length 3.6 3.2 3.6 3.1 3.2CLI – Clasper �nner length 6.8 6.3 6.3 6.3 6.3CLB – Clasper base w�dth 1.3 1.5 1.6 1.5 1.5

stra�ght, not vert�cal, d�rected well anterodorsally from bottom to top, moderately concave near free rear t�p (less so �n paratype); free rear t�p narrow basally (otherw�se moderate); inner margin of fin almost straight; insertion of base extremely well forward of pelvic-fin origin, well poster�or (by about an eye d�ameter) to free rear t�p of pectoral fin; fin-spine origin well posterior to pectoral-fin insertion; spine base broad, exposed anteriorly well below junction of spine and soft portion of fin (almost three-quarters of spine exposed); soft portion of fin connected sl�ghtly above m�d-po�nt of total sp�ne length; sp�ne taper�ng only sl�ghtly d�stally, anter�or marg�n almost stra�ght; exposed port�on more upr�ght, subequal in length to exposed portion of second dorsal-fin spine; pre-first dorsal length 3.63 (3.70) times in TL; first dorsal-fin length 1.89 (1.90) times its height, 1.14 (1.19) times second dorsal-fin length; first dorsal-fin height

1.72 (1.64) times second dorsal-fin height; exposed first dorsal spine length 0.64 (0.68) times height of fin. Second dorsal fin small, strongly raked; anterior margin moderately convex, apex narrowly rounded; poster�or marg�n very deeply concave, max�mum concav�ty almost near m�d-po�nt of marg�n, upper port�on d�rected dorsoposter�orly strongly from bottom to top; free rear t�p relat�vely short (or moderate), �nner marg�n length 0.96 (0.92) times fin height; second dorsal-fin length 2.84 (2.61) t�mes �ts he�ght; sp�ne length 1.03 (1.00) �n he�ght of fin; fin-spine origin well behind free rear tip of pelvic fin, exposed well below level of junction with spine and soft portion of fin, proximal part of spine base close to dorsal profile of body; second spine moderately broad based, slender, not sharply po�nted d�stally (poss�bly w�th m�nor ap�cal damage �n both types), not taper�ng rap�dly just above po�nt of exposure, sp�ne t�p extend�ng just


behind (otherwise over) level of insertion of fin, much more strongly raked than first dorsal spine; interdorsal space 0.93 (0.88) �n pre-pectoral length, 1.16 (1.09) �n pre-first dorsal length; moderate interdorsal groove. Pectoral fin small, anterior margin weakly convex; inner marg�n stra�ght to sl�ghtly convex (otherw�se moderately convex), length 7.1 (6.6)% TL; apex narrowly rounded, not lobe-l�ke or falcate; poster�or marg�n weakly concave (fin weakly falcate with concave posterior margin in paratype), free rear t�p broadly rounded; base very short, 2.67 (2.76) in length of anterior margin. Pelvic fins small, anter�or and poster�or marg�ns almost stra�ght, apex broadly rounded, free rear t�p acute. Caudal peduncle long, tapering slightly to caudal fin; subcircular in cross-sect�on anter�orly, broadly sem�c�rcular poster�orly, ventral groove very well developed; lateral keels well developed, or�g�nat�ng under �nsert�on of second dorsal fin, terminating about half or less than an eye diameter behind caudal-fin insertion; pelvic-caudal space 0.81 (0.84) �n pectoral–pelv�c space, 0.82 (0.83) �n prepectoral length; dorsal–caudal space 2.08 (2.11) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal pit rudimentary. Caudal fin short, dorsal caudal margin 1.12 (1.09) �n head length; length of lower caudal lobe 1.81 (1.75) �n upper lobe length; upper poster�or lobe strongly convex, base of lobe broad; lower lobe apex narrowly rounded (more angular �n paratype). Vertebral centra 120 (114 �n paratype), monospondylous 44 (42), precaudal 92 (88) and caudal 28 (26). Teeth �n upper jaw (of paratype, CSIRO CA 3297) 13+14=27, lower jaw 11+11=22.

COLOUR.— In preservat�ve: dorsal half pale yellow�sh grey, off wh�te ventrally; pale and dark areas on head and trunk not strongly demarcated, tones merg�ng gradually. First dorsal-fin base off white, strongly demarcated from body and main portion of fin; soft portion of fin uniformly light grey, margin paler, no dark areas at fin apex; second dorsal fin similar to first dorsal, base also off white, strongly demarcated from rest of fin; both

Figure 7. Cusps of the flank denticles of Squalus altipinnis sp. nov. paratype (CSIRO CA 3297, adult male 589 mm TL). F�eld of v�ew w�dth 1.4 mm.

dorsal-fin spines almost uniformly dark grey (slightly darker on anterior edge of first dorsal fin in holotype), much darker than base of soft portion of fin. Caudal fin l�ght grey, paler along poster�or marg�n �n both lobes; no dark areas on fin. Pectoral and pelvic fins similar to first dorsal fin, uniformly light grey with a narrow, pale posterior margin; fin bases off white on ventral surface.

SIZE.— Presumably a small spurdog, known only from two adult male spec�mens, 586 and 589 mm TL.

DISTRIBUTION.— Cont�nental slope off northwestern Austral�a �n the v�c�n�ty of the Rowley Shoals (ca. 17– 18° S) �n about 300 m depth.

ETYMOLOGY.— Der�ved from the Lat�n comb�nat�on of altus (h�gh) and pinna (fin) in allusion to the upright dorsal fin, typical of members of this subgroup.

VERNACULAR.— Western Highfin Spurdog.

REMARKS.— Squalus altipinnis d�ffers from S. albifrons, in having a smaller caudal fin (dorsal caudal marg�n 19.6–19.9 vs. 21.3–23.6% TL �n S. albifrons; preventral caudal marg�n 10.8–11.4 vs. 11.2–12.5% TL; caudal fork length 8.4–8.5 vs. 9.4–10.5% TL), smaller pectoral fin (anterior margin 13.9–14.1 vs. 14.2–15.8% TL; poster�or marg�n 10.2–10.4 vs. 10.8–11.7% TL)

Figure 8. Lateral view of the dorsal fins of: Squalus altipinnis sp. nov. holotype (CSIRO CA 4111, adult male 586 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B

48

and longer dorsal–caudal space (11.4–11.7 vs. 9.4–11.1% TL). Some other rat�os appear to be useful for distinguishing these species: pre-second dorsal-fin length 4.32–4.36 (3.71–4.28) times pectoral-fin anterior margin, 3.07–3.08 (2.48–2.84) t�mes dorsal caudal marg�n, 4.16–4.22 (3.18–3.89) t�mes upper post ventral marg�n; mouth w�dth 3.11–3.26 (3.29–4.08) t�mes upper lab�al furrow; head length 2.90–2.96 (2.52–2.73) t�mes mouth w�dth; pre-vent length about 2.48 (2.00–2.24) t�mes dorsal caudal marg�n; dorsal caudal marg�n 1.70–1.72 (1.94–2.48) t�mes dorsal–caudal space.

Squalus altipinnis differs from a third new highfin spurdog descr�bed �n th�s paper, (�.e. S. sp. A: sensu Last and Stevens, 1994), in having a smaller caudal fin (dorsal caudal marg�n 19.6–19.9 vs. 23.3–24.0% TL �n S. sp. A; preventral caudal marg�n 10.8–11.4 vs. 11.5–11.9% TL; precaudal length 79.9–80 vs. 74.9–76.1% TL), smaller anterior pectoral-fin margin (13.9–14.1 vs. 14.6–16.4% TL), and longer dorsal caudal space (11.4–11.7 vs. 9.5–10.8% TL). Several other rat�os appear to be useful for d�st�ngu�sh�ng these spec�es: pre-second dorsal length 4.32–4.36 (vs. 3.60–3.98 �n S. sp. A) times pectoral-fin anter�or marg�n, 3.07–3.08 (vs. 2.39–2.57) t�mes dorsal caudal marg�n, 4.16–4.22 (vs. 3.16–3.33) t�mes upper post ventral marg�n; mouth w�dth 3.11–3.26 (vs. 3.45–4.28) t�mes upper lab�al furrow; head length 2.90–2.96 (vs. 2.63–2.79) t�mes mouth w�dth; pre-vent length about 2.48 (vs. 1.88–2.00) t�mes dorsal caudal marg�n; dorsal caudal marg�n 1.70–1.72 (vs. 2.21–2.46) t�mes dorsal–caudal space, 1.75–1.81 (vs. 2.01–2.09) t�mes pre-ventral caudal marg�n; head length 4.41–4.72 (vs. 3.77–4.26) t�mes eye length; 1.09–1.12 (vs. 0.85–0.89) t�mes dorsal caudal margin; and first dorsal-fin length 1.89–1.90 (vs. 1.50–1.68) t�mes �ts he�ght.

Squalus notocaudatus sp. nov.


Squalus sp. A: Last and Stevens, 1994, Sharks and Rays of Australia, pp 48, 95, figs 19, 8.31, pl. 6; Compagno, Dando and Fowler, 2005, Sharks of the world, p 78–79, figs, pl. 2. Holotype. CSIRO H 1368–02, �mmature male 619 mm TL, east of Flinders Reef, Queensland, 17°27′ S, 149° 46′ E, 348 m, 3 Dec 1985.Paratypes. 3 spec�mens. CSIRO H 1322–01, female 393 mm TL, east of Flinders Reef, Queensland, 17°28′ S, 149°41′ E, 402 m; CSIRO H 1323–01, immature male 368 mm TL, Queensland Plateau, east of Fl�nders Reef, Queensland, 17°32′ S, 149°34′ E, 454 m; CSIRO H 1321–01, female 495 mm TL, Capr�corn Channel, east of Rockhampton, Queensland, 22°52′ S, 152°42′ E, 282 m.

DIAGNOSIS.— A moderate-s�zed or large spec�es of Squalus of the ‘highfin megalops group’ with the follow�ng comb�nat�on of characters: abdomen depth 8.6–

11.1% TL; pre-vent length 45.3–46.5% TL, 1.9–2.0 t�mes dorsal caudal marg�n; pre-second dorsal length 3.6–4.0 times pectoral-fin anterior margin, 2.4–2.6 times dorsal caudal marg�n; head w�dth 1.4–1.6 t�mes abdomen w�dth; preoral length 2.8–3.2 t�mes hor�zontal prenar�al length, 8.4–9.5% TL; head length 3.8–4.3 t�mes eye length; mouth w�dth 3.4–4.3 t�mes length of upper lab�al furrow; �nterorb�tal w�dth 1.4–1.5 t�mes hor�zontal preorb�tal length; fifth gill slit height 1.9–2.1% TL; anterior nasal flap strongly bifurcate; first dorsal fin upright, upper poster�or marg�n almost vert�cal, greatest concav�ty closer to free rear tip than fin apex; posterior margin of second dorsal fin deeply concave; second dorsal-fin spine with a broad base; pectoral fin of adult weakly falcate, anter�or marg�n long, 14.6–16.4% TL; both dorsal and ventral surfaces pale; dorsal fins pale with obvious dark tips; first dorsal-fin spine and base of soft portion of dorsal fin both pale; caudal fin with broad white posterior marg�n; prom�nent dark, d�agonal subterm�nal streak parallel to the upper posterior margin; flank denticles strongly tr�cusp�d; 47–49 monospondylous centra, 94–97 precaudal centra, 123–127 total centra; largest spec�men �mmature at 62 cm TL.

DESCRIPTION.— Body fus�form, slender, nape not prominently humped; deepest near first dorsal-fin spine, max�mum depth 1.06 (0.94–1.06 �n paratypes) t�mes w�dth, trunk depth 1.08 (0.83–0.95) t�mes abdomen depth; head short 19.7 (20.4–21.1)% TL; caudal peduncle slender, 25.6 (25.2–27.1)% TL. Head rather broad, much w�der than trunk, w�dth 1.34 (1.24–1.41) t�mes trunk w�dth, 1.46 (1.42–1.60) t�mes abdomen w�dth, depressed forward of sp�racles, becom�ng subtr�angular towards pectoral-fin origin, length 2.36 (2.18–2.22) in pre-vent length; he�ght 0.74 (0.71–0.78) t�mes w�dth. Snout short, narrowly tr�angular �n lateral v�ew, apex bluntly po�nted, lateral prenar�al marg�n sl�ghtly angular; narrowly rounded �n dorsal v�ew, d�st�nct prenar�al notch (not ev�dent �n paratype CSIRO H 1321–01), hor�zontal length 1.19 (1.10–1.31) t�mes eye length, 0.68 (0.68–0.72) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.87 (2.83–3.17) t�mes �n preoral length. Eye narrowly oval, s�ze moderate, length 4.23 (3.77–4.26) �n head length, 2.81 (2.32–3.45) t�mes depth; strongly notched poster�orly, notch extend�ng as a well-developed (less developed �n two smallest paratypes, CSIRO H 1322–01 and CSIRO H 1323–01) furrow to anteroventral marg�n of sp�racle. Sp�racle moderate, subtr�angular (var�ably closed �n paratypes); narrow lobe-l�ke fold on poster�or marg�n, �n paratypes more strongly produced w�th a convex anter�or marg�n; greatest d�ameter 4.03 (3.60–4.44) �n eye length. G�ll open�ngs almost upr�ght (variable in paratypes), first four subequal in size, fifth usually slightly longer, height of fifth slit 1.9 (2.0–2.1)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.15 (1.19–1.26) �n preoral length; upper lab�al furrows about tw�ce or less length of lower furrows; prom�nent postoral groove, subequal �n length to upper lab�al furrows, extend�ng posterolaterally from angle of


Figure 9. Lateral v�ew of: Squalus notocaudatus sp. nov. holotype (CSIRO H 1368–02, �mmature male 619 mm TL; �mage reversed).

Figure 10. Ventral v�ew of the head of Squalus notocaudatus sp. nov. holotype (CSIRO H 1368–02, �mmature male 619 mm TL).

jaws; two ser�es of funct�onal teeth �n upper jaw, two ser�es �n lower. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Nostr�ls small, almost transverse; anterior nasal flap strongly bifurcate, upper lobe largest, broadest; posterior lobe narrow, finger-like; internarial space 1.91 (1.92–2.25) �n preoral length, 2.47 (2.19–2.56) t�mes nostr�l length. Dermal dent�cles (based on paratypes CSIRO H 1321–01 and CSIRO H 1322–01) on flank very small, strongly tricuspidate with pronounced med�an r�dge; med�an r�dge commenc�ng well anter�or to rest of crown, w�th a mes�al furrow develop�ng anter�orly and converg�ng towards poster�or t�p of crown; lateral cusps well to very well developed, acute poster�orly, r�dge anter�or, not d�stal or extend�ng toward t�p of cusp; weakly or not �mbr�cated, more regular on snout t�p. F�rst dorsal fin elevated, upright, narrowly rounded apically; anter�or marg�n strongly convex; upper poster�or marg�n

almost stra�ght, subvert�cal, not d�rected anterodorsally from bottom to top (sl�ghtly anterodorsally �n paratype CSIRO H 1321–01), moderately concave near free rear t�p; free rear t�p moderately short (less well developed �n some paratypes), moderately th�ck basally; �nner marg�n of fin almost straight; insertion of base extremely well forward of pelvic-fin origin, slightly posterior to free rear tip of pectoral fin; fin-spine origin more-or-less above pectoral-fin insertion; spine base broad, exposed anteriorly just below junction of spine and fin; soft portion of fin connected near mid-point of total spine length (skin cover�ng smallest paratypes extend�ng toward sp�ne apex, �n paratype CSIRO H 1323–01 exposed he�ght of sp�ne equ�valent to he�ght of sp�racle); sp�ne taper�ng weakly d�stally (var�able �n paratypes), anter�or marg�n weakly convex (more strongly convex �n paratype CSIRO H 1321–01); exposed port�on raked sl�ghtly, shorter �n length to exposed portion of second dorsal-fin spine; pre-first dorsal length 3.81 (3.64–3.73) times in TL; first

50

dorsal-fin length 1.68 (1.50–1.66) times its height, 1.13 (1.10–1.19) times second dorsal-fin length; first dorsal-fin height 1.74 (1.71–1.77) times second dorsal-fin height; exposed first dorsal spine length 0.55 (0.40–0.47) times height of fin. Second dorsal fin of moderate size, strongly raked; anter�or marg�n strongly convex, apex narrowly rounded; poster�or marg�n deeply concave, angle about 90º or sl�ghtly more, max�mum concav�ty almost near m�d-po�nt of marg�n, upper port�on d�rected dorsoposter�orly strongly from bottom to top; free rear t�p elongate, �nner marg�n length 0.98 (0.86–1.01) t�mes fin height; second dorsal-fin length 2.59 (2.22–2.65) t�mes �ts he�ght; sp�ne length 1.20 (0.90–0.99) �n he�ght of fin; fin-spine origin well behind free rear tip of pelvic fin, exposed just below level of junction with spine and soft portion of fin; second spine moderately broad based, slender, part�ally covered anter�orly by sk�n �n two smallest paratypes), sharply po�nted d�stally, taper�ng evenly from po�nt of exposure, sp�ne t�p extend�ng behind level of insertion of fin (over in paratype CSIRO H 1323–01), much more strongly raked than first spine; �nterdorsal space 0.78 (0.87–0.89) �n length from snout tip to pectoral-fin origin, 1.04 (1.11–1.17) in pre-first dorsal length; moderate �nterdorsal groove (weaker �n paratypes). Pectoral fin well developed, anterior margin weakly convex; �nner marg�n moderately convex, length 7.3 (7.5–7.8)% TL; apex narrowly rounded, lobe-l�ke, somewhat falcate (less pronounced �n smallest paratypes); poster�or marg�n strongly concave, free rear t�p bluntly angular; base very short, 3.00 (2.67–3.26) �n length of anterior margin. Pelvic fins small, anterior and posterior marg�ns almost stra�ght, apex broadly rounded, free rear t�p acute. Caudal peduncle long, taper�ng sl�ghtly to caudal fin; subcircular in cross-section anteriorly, almost sem�c�rcular poster�orly, ventral groove very well developed; lateral keels poorly developed, or�g�nat�ng anter�or to (under �n some paratypes) �nsert�on of second dorsal fin, terminating about half an eye diameter behind caudal-fin insertion; pelvic–caudal space 0.85 (0.73–0.83)

Figure 11. Cusps of the flank denticles of Squalus notocaudatus sp. nov. paratype (CSIRO H 1322–01, female 393 mm TL). F�eld of v�ew w�dth 0.8 mm.

�n pectoral–pelv�c space, 0.76 (0.78–0.83) �n prepectoral length; dorsal–caudal space 2.66 (2.15–2.40) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal pit rudimentary. Caudal fin relatively elongate, dorsal caudal marg�n 0.85 (0.85–0.89) �n head length; length of lower caudal lobe 2.01 (2.02–2.09) �n upper lobe length; upper poster�or lobe moderately convex; lower lobe apex somewhat angular (more rounded �n paratypes). Vertebral centra 127 (123–125 �n 3 paratypes), monospondylous 49 (47–49), precaudal 97 (94–96) and caudal 30 (29). Teeth �n upper jaw (of paratype, CSIRO H 1321–01) 14+13=27, lower jaw 12+11=23.

COLOUR.— When fresh (based on holotype): un�form pale grey dorsally; not significantly paler on ventral surface, dorsal and ventral tonal areas merg�ng gradually, not sharply demarcated. First dorsal fin pale grey, anterior basal half wh�t�sh, narrow black edge to upper anter�or margin; second-dorsal fin pale greyish, slightly paler through base and free rear t�p, th�n black marg�n along anterior, apical and posterior margins; first dorsal-fin spine pale; second dorsal-fin spine dusky, anterior margin darkest. Caudal fin upper lobe with thin black anterior fr�nge; poster�or marg�n broadly wh�te, rema�n�ng upper lobe pale grey; ventral lobe pale; prom�nent black�sh, d�agonal, bar–l�ke mark�ng extend�ng from near anter�or

Figure 12. Lateral view of the dorsal fins of: Squalus notocaudatus sp. nov. holotype (CSIRO H 1368–02, immature male 619 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B


Table 2. Proport�onal d�mens�ons as percentages of total length for the holotype (CSIRO H 1368–02) and ranges for the 3 paratypes of Squalus notocaudatus sp. nov.

S. notocaudatus sp. nov. Holotype Paratypes

M�n. Max.TL — Total length 619 368 495PCL — Precaudal length 75.9 74.9 76.1PD2 — Pre-second dorsal length 59.8 57.4 58.2PD1 — Pre-first dorsal length 26.2 26.8 27.5SVL — Pre-vent length 46.5 45.3 46.1PP2 — Prepelv�c length 44.9 44.2 44.8PP1 — Prepectoral length 19.6 20.6 21.2HDL — Head length 19.7 20.4 21.1PG1 — Prebranch�al length 17.0 17.6 18.3PSP — Presp�racular length 11.3 12.1 12.2POB — Preorb�tal length 6.4 6.9 7.0PRN — Prenar�al length 4.3 4.5 5.0POR — Preoral length 8.4 9.3 9.5INLF — Inner nostr�l-lab�al furrow space 4.5 4.6 4.9MOW — Mouth w�dth 7.3 7.3 8.0ULA — Lab�al furrow length 2.0 1.8 2.1INW — Internar�al space 4.4 4.2 4.8INO — Interorb�tal space 8.1 8.8 9.2EYL — Eye length 4.7 4.8 5.5EYH — Eye he�ght 1.7 1.6 2.1SPL — Sp�racle length 1.2 1.2 1.4GS1 — F�rst g�ll-sl�t he�ght 1.8 2.0 2.1GS5 — F�fth g�ll-sl�t he�ght 1.9 2.0 2.1IDS — Interdorsal space 25.2 23.4 24.2DCS — Dorsal-caudal space 9.5 10.1 10.8PPS — Pectoral-pelv�c space 21.6 19.1 20.9PCA — Pelv�c-caudal space 25.6 25.2 27.1D1L — F�rst dorsal length 13.7 13.6 14.1D1A — F�rst dorsal anter�or marg�n 11.9 12.9 13.7D1B — F�rst dorsal base length 8.5 7.9 9.0D1H — F�rst dorsal he�ght 8.2 8.3 9.4D1I — F�rst dorsal �nner marg�n 5.6 5.0 5.7D1P — F�rst dorsal poster�or marg�n 9.1 8.8 10.5D1ES — F�rst dorsal sp�ne length 4.5 3.6 4.4D1BS — F�rst dorsal sp�ne base w�dth 1.0 0.7 0.9D2L — Second dorsal length 12.1 11.8 12.5D2A — Second dorsal anter�or marg�n 10.2 10.5 11.5D2B — Second dorsal base length 7.6 7.2 8.2D2H — Second dorsal he�ght 4.7 4.7 5.3D2I — Second dorsal �nner marg�n 4.6 4.5 4.7D2P — Second dorsal poster�or marg�n 5.4 4.7 5.7D2ES — Second dorsal sp�ne length 5.6 4.2 5.3D2BS — Second dorsal sp�ne base w�dth 0.9 0.8 0.8

52

Table 1. cont’d

S. notocaudatus sp. nov. Holotype Paratypes

M�n. Max.P1A — Pectoral anter�or marg�n 16.4 14.6 16.0P1I — Pectoral �nner marg�n 7.3 7.5 7.8P1B — Pectoral base length 5.5 4.9 5.5P1P — Pectoral poster�or marg�n 10.6 9.9 11.4P2L — Pelv�c length 9.4 9.1 9.6P2H — Pelv�c he�ght 4.4 4.3 4.8P2I — Pelv�c �nner marg�n 4.7 4.4 5.1CDM — Dorsal caudal marg�n 23.3 23.4 24.0CPV — Preventral caudal marg�n 11.6 11.5 11.9CPU — Upper postventral caudal marg�n 17.9 17.5 18.2CPL — Lower postventral caudal marg�n 4.1 3.7 4.7CFW — Caudal fork w�dth 6.3 6.5 6.9CFL — Caudal fork length 9.1 8.6 9.9HANW — Head w�dth at nostr�ls 6.7 6.7 7.1HAMW — Head w�dth at mouth 10.9 10.9 11.5HDW — Head w�dth 11.8 12.7 12.7TRW — Trunk w�dth 8.9 9.0 10.2ABW — Abdomen w�dth 8.1 7.9 9.0TAW — Ta�l w�dth 5.7 5.5 6.1CPW — Caudal peduncle w�dth 2.1 2.5 2.6HDH — Head he�ght 8.7 8.9 10.0TRH — Trunk he�ght 9.4 9.2 10.2ABH — Abdomen he�ght 8.6 10.3 11.1TAH — Ta�l he�ght 5.9 6.1 7.1CPH — Caudal peduncle he�ght 2.4 2.5 2.6CLO — Clasper outer length 1.4 0.6 0.6CLI — Clasper �nner length 3.2 2.7 2.7CLB — Clasper base w�dth 0.7 0.5 0.5

marg�n of lower lobe towards caudal fork and on to sub-basal port�on of upper lobe. In preservat�ve (based on holotype): holotype more un�formly toned than �n fresh condition, somewhat yellowish brown; fins with dark markings less evident; pectoral and pelvic fins uniform (possibly with indistinct pale edges on both fins); tip of snout w�th �rregular black�sh mark�ng. In small paratype (CSIRO H 1323–01), black snout mark�ng absent, dark areas on fins more strongly developed than on fresh holotype; free rear tip of first dorsal fin whitish, distinctly paler than non-basal portion of fin; blackish area on second dorsal fin more pronounced, free rear tip pale; dark anterior marking on upper lobe of caudal fin broader than on holotype; dark streak through lower half of fin well defined, strongly contrasted with broad whitish marg�n at caudal fork.

SIZE.— Known from l�m�ted mater�al. Largest spec�men �s the 619 mm TL, �mmature male holotype (CSIRO H 1368–02). The 3 paratypes range from 368 to 495 mm TL.

DISTRIBUTION.— Based on a few spec�mens collected off central Queensland near Fl�nders Reef (ca. 17° S) and off Rockhampton (ca. 23° S) �n 225–454 m depth.

ETYMOLOGY.— Der�ved from a comb�nat�on of the Lat�n nota (mark) and cauda (ta�l) w�th reference to the dark bar on the caudal fin.

VERNACULAR.— Barta�l Spurdog.

REMARKS.— Squalus notocaudatus �s most s�m�lar morphometr�cally to S. albifrons, but d�ffers �n the


follow�ng characters: head length 3.8–4.3 (4.3–4.9 �n S. albifrons) t�mes eye length; �nterorb�tal space 1.39–1.47 (1.48–1.73) t�mes hor�zontal preorb�tal length; dorsal caudal marg�n 2.0–2.1 (1.9–2.0) t�mes pre-ventral caudal marg�n; pre-vent length 1.9–2.0 (2.0–2.4) t�mes length of dorsal caudal margin; first dorsal-fin length 1.5–1.7 (1.7–1.8) times its height; fifth gill-slit length 1.9–2.1 (2.1–2.5)% TL; pelvic-fin inner margin length 4.4–5.1 (5.0–6.0)% TL; upper lab�al furrow 1.8–2.1 (2.1–2.4)% TL; and caudal peduncle w�dth 2.1–2.6 (2.8–3.2)% TL. Squalus notocaudatus �s much paler coloured on the dorsal surface than S. albifrons, and the s�de of the head does not d�splay sharp tonal d�fferences. It also has more vertebrae (94–97 vs. 89–93 precaudal centra), and the posterior margin of the first dorsal fin of S. notocaudatus, wh�ch �s or�ented almost vert�cally (rather than be�ng d�rected posteroventrally), �s more strongly concave w�th the deepest part of the concav�ty s�tuated more basally (i.e. closer to the free rear tip than the fin apex rather than about midway between the free rear tip and the fin marg�n).

Squalus notocaudatus d�ffers from S. altipinnis, �n hav�ng a much larger caudal fin (dorsal caudal margin 23.3–24.0 vs. 19.6–19.9% TL �n S. altipinnis; preventral caudal marg�n 11.5–11.9 vs. 10.8–11.4% TL; precaudal length 74.9–76.1 vs. 79.9–80% TL), larger anter�or pectoral-fin margin (14.6–16.4 vs. 13.9–14.1% TL), and shorter dorsal–caudal space (9.5–10.8 vs. 11.4–11.7% TL). Some other rat�os appears to be useful for d�st�ngu�sh�ng these spec�es: pre-second dorsal length 3.60–3.98 (vs. 4.32–4.36 �n S. altipinnis) times pectoral-fin anterior marg�n, 2.39–2.57 (vs. 3.07–3.08) t�mes dorsal caudal marg�n, 3.16–3.33 (vs. 4.16–4.22) t�mes upper post ventral marg�n; mouth w�dth 3.45–4.28 (vs. 3.11–3.26) t�mes upper lab�al furrow; head length 2.63–2.79 (vs. 2.90–2.96) t�mes mouth w�dth; pre-vent length 1.88–2.00 (vs. 2.48) t�mes dorsal caudal marg�n; dorsal caudal marg�n 2.21–2.46 (vs. 1.70–1.72) t�mes dorsal–caudal space, 2.01–2.09 (vs. 1.75–1.81) t�mes pre-ventral caudal marg�n; head length 3.77–4.26 (vs. 4.41–4.72) t�mes eye length; 0.85–0.89 (vs. 1.09–1.12) t�mes dorsal caudal margin; and first dorsal-fin length 1.50–1.68 (vs. 1.89–1.90) t�mes �ts he�ght.

ACKNOWLEDGEMENTS

The new species were first identified in a study of the Austral�an chondr�chthyan fauna (Last and Stevens, 1994) that was jo�ntly funded by the CSIRO and the F�sher�es Research and Development Corporat�on (FRDC). The authors would l�ke to thank Alasta�r Graham for assembl�ng data on Austral�an mater�als, Lou�se Conboy for prepar�ng �mages of the types, T�m Founta�n for prepar�ng X-rays and obta�n�ng mer�st�c �nformat�on, Tony Rees for ass�st�ng w�th preparat�on of SEMs, Matt Edmunds for h�s extens�ve work on th�s group, Dan�el Gledh�ll for prepar�ng a l�terature summary, Gordon Yearsley and John Pogonosk� for other techn�cal support and ed�tor�al ass�stance. Thanks also go to Jeff Johnson (QM) for h�s techn�cal ass�stance and to Ken Graham for collect�on of some mater�al.

REFERENCES


Graham, K.J. (2005) D�str�but�on, populat�on structure and b�olog�cal aspects of Squalus spp. (Chondr�chthyes: Squal�formes) from New South Wales and adjacent Austral�an waters. Marine and Freshwater Research, 56, 405–416.


Last, P.R., Wh�te, W.T., Pogonosk�, J.J., Gledh�ll, D.C., Ward, B. and Yearsley, G.K. (2007) Part 1 — Appl�cat�on of a rap�d taxonom�c approach to the genus Squalus, p. 1–10. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.


Part 6 — Description of Squalus chloroculus sp. nov., a new spurdog from southern Australia, and the resurrection of S. montalbani Whitley

Peter R. Last1, W�ll�am T. Wh�te1 and H�royuk� Motomura2

1CSIRO Mar�ne & Atmospher�c Research, GPO Box 1538, Hobart, TAS, 7001, AUSTRALIA2The Kagosh�ma Un�vers�ty Museum, 1–21–30 Kor�moto, Kagosh�ma 890–0065, JAPAN

ABSTRACT.— A new spurdog, Squalus chloroculus sp. nov., �s descr�bed based on spec�mens from the upper continental slope off southern Australia. This species and a closely related Indo–West Pacific species, S. montalbani Whitley, have been consistently confused with each other and misidentified as a western North Pacific spurdog, S. mitsukurii Jordan and Snyder. In th�s paper, the new spec�es �s descr�bed and Squalus montalbani �s resurrected and red�agnosed based on mater�al from the Ph�l�pp�nes, Indones�a, and tropical and warm temperate Australia. Intraspecific variation between populations of S. montalbani across these reg�ons �s d�scussed. These spec�es are compared to S. mitsukurii and belong to a subgroup of Squalus, the ‘m�tsukur�� group’, whose members have a moderately elongate snout, stocky body, and a dark caudal bar on the posterior notch of the caudal fin. These spurdogs are very similar morphologically but can be distinguished using molecular techniques and through a combination of caudal-fin coloration, meristics, and morphometrics of the head, trunk and fins.

Key words. Squalo�dea – Squal�dae – Squalus chloroculus – new spec�es – Squalus montalbani – resurrected spec�es – southern Austral�a


INTRODUCTION

Squalus mitsukurii Jordan and Snyder �n Jordan and Fowler, 1903, wh�ch has long been cons�dered to be a very wide ranging dogfish in temperate and tropical oceans (Compagno, 1984), �s now thought to be a spec�es complex (Last and Stevens, 1994; Compagno et al., 2005). Members of th�s complex, referred to here as the ‘m�tsukur�� group’, are character�sed by the�r large relat�ve size, a dark caudal bar, low dorsal-fin spines, and a small, raked first dorsal fin. An Australian form referable to S. mitsukurii was reported by Last and Stevens (1994) from the temperate and trop�cal cont�nental slope of Austral�a between Townsv�lle (Queensland) and Shark Bay (Western Austral�a). Molecular stud�es (see Ward et al., 2007, Part 12 of th�s �ssue) of Austral�an members of th�s complex revealed the ex�stence of two spec�es. One of these spurdogs has a temperate d�str�but�on off southern Austral�a, whereas the other spec�es occurs pr�mar�ly �n subtrop�cal and trop�cal lat�tudes off both eastern and western Austral�a. These spec�es are very s�m�lar morpholog�cally to each other and to S. mitsukurii.

Market surveys at var�ous land�ng s�tes �n the Ph�l�pp�nes (late 1990’s) and eastern Indones�a (Apr�l 2001 to March 2006) produced a w�de var�ety of sharks, rays and ch�maeras, �nclud�ng many squalo�ds, for research. Th�s mater�al �ncluded mult�ple spec�es of Squalus, one

of which appeared to be conspecific with an Australian member of the ‘m�tsukur�� group’. A Ph�l�pp�ne spec�es, which was provisionally identified as S. cf. mitsukurii, was thought to be non-conspecific with S. mitsukurii (Compagno et al., 2005). Sm�th and Radcl�ffe �n Sm�th (1912) descr�bed Squalus philippinus (a jun�or homonym of S. philippinus Shaw, 1804 = Heterodontus portusjacksoni (Meyer, 1793)) from the west coast of Luzon Island, Ph�l�pp�nes. Th�s spec�es, wh�ch was later renamed Squalus montalbani by Wh�tley (1931), was cons�dered to be a l�kely jun�or synonym of S. mitsukurii (Compagno, 1984). In the follow�ng paper, the temperate Austral�an spurdog �s descr�bed as a new spec�es and Squalus montalbani �s resurrected and red�agnosed. These spec�es are compared to the types and other mater�al, from near the type local�ty (off Japan), of Squalus mitsukurii.

METHODS

Methods follow those outl�ned �n Part 1 of th�s �ssue (Last et al., 2007). D�agnoses are prov�ded for S. montalbani and the new spec�es; mean values for d�agnost�c rat�os and counts are g�ven �n parentheses after the�r ranges. In the new spec�es descr�pt�on, morphometr�c and meristic values for the holotype are given first followed �n parentheses by the ranges of the measured paratypes.

56

Table 1. Proport�onal d�mens�ons as percentages of total length for the holotypes of Squalus montalbani (USNM 70256) and Squalus mitsukurii (SU 12793) and ranges for add�t�onal mater�al measured.

S. montalbani S. mitsukurii n = 14 n = 4

Holotype M�n. Max. Holotype M�n. Max.TL – Total length 311 520 843 719 266 855PCL – Precaudal length 76.5 77.1 79.9 76.6 78.2 79.0PD2 – Pre-second dorsal length 57.6 59.1 62.8 59.8 58.6 61.2PD1 – Pre-first dorsal length 30.7 26.5 30.0 30.9 28.5 32.3SVL – Pre-vent length 47.2 48.6 52.7 51.5 48.9 52.2PP2 – Prepelv�c length 43.8 47.2 50.8 48.5 47.4 50.1PP1 – Prepectoral length 22.8 20.8 22.9 23.3 19.9 23.9HDL – Head length 23.2 21.3 23.4 23.4 20.9 23.5PG1 – Prebranch�al length 20.3 17.6 19.7 19.5 18.0 20.1PSP – Presp�racular length 13.4 11.3 13.6 12.8 12.1 13.3POB – Preorb�tal length 7.1 6.7 8.1 7.5 7.3 7.9PRN – Prenar�al length 4.6 4.0 5.3 5.5 5.0 5.4POR – Preoral length 10.1 8.7 10.6 10.8 9.4 10.6INLF – Inner nostr�l-lab�al furrow space 5.3 4.2 4.9 4.4 4.2 4.7MOW – Mouth w�dth 6.5 7.1 8.5 6.2 6.3 7.5ULA – Lab�al furrow length 2.5 1.9 2.5 2.4 2.1 2.5INW – Internar�al space 5.2 4.1 5.0 4.8 4.0 4.9INO – Interorb�tal space 8.7 7.5 9.6 8.1 7.9 8.4EYL – Eye length 4.3 3.8 5.2 3.4 3.8 4.7EYH – Eye he�ght 1.6 1.5 2.3 1.3 1.8 2.5SPL – Sp�racle length 1.5 1.3 1.9 1.2 1.2 1.5GS1 – F�rst g�ll-sl�t he�ght 1.6 1.7 2.5 1.9 1.6 1.7GS5 – F�fth g�ll-sl�t he�ght 2.0 1.7 2.4 2.1 1.8 2.0IDS – Interdorsal space 21.7 22.1 25.9 21.3 18.7 25.2DCS – Dorsal-caudal space 11.2 9.7 11.2 9.8 9.9 11.2PPS – Pectoral-pelv�c space 18.9 19.8 25.4 22.5 21.3 24.5PCA – Pelv�c-caudal space 26.3 22.9 26.0 22.7 22.3 27.4D1L – F�rst dorsal length 14.2 12.8 15.9 14.5 12.5 15.7D1A – F�rst dorsal anter�or marg�n 12.4 10.4 12.6 12.0 10.5 11.1D1B – F�rst dorsal base length 8.3 7.6 9.8 8.3 7.8 7.8D1H – F�rst dorsal he�ght 6.1 5.7 7.4 8.5 4.5 8.3D1I – F�rst dorsal �nner marg�n 6.2 4.9 6.8 6.3 4.9 6.4D1P – F�rst dorsal poster�or marg�n 6.7 6.9 8.8 9.7 4.6 7.9D1ES – F�rst dorsal sp�ne length 2.1 2.2 4.0 3.3 3.5 4.8D1BS – F�rst dorsal sp�ne base w�dth 0.7 0.5 0.6 0.8 0.6 0.8D2L – Second dorsal length 13.4 11.1 13.9 12.7 12.0 13.9D2A – Second dorsal anter�or marg�n 10.2 8.5 10.8 10.2 10.4 10.7D2B – Second dorsal base length 8.1 7.0 8.5 7.2 8.0 9.2D2H – Second dorsal he�ght 4.2 3.4 4.6 4.5 3.0 4.6D2I – Second dorsal �nner marg�n 5.9 4.0 5.7 5.1 4.2 5.4D2P – Second dorsal poster�or marg�n 5.2 4.5 6.3 5.2 4.1 4.4D2ES – Second dorsal sp�ne length 3.6 2.0 3.9 3.8 3.8 5.0D2BS – Second dorsal sp�ne base w�dth 0.8 0.5 0.6 0.7 0.7 0.9


Table 1. cont’d.

S. montalbani S. mitsukurii n = 14 n = 4

Holotype M�n. Max. Holotype M�n. Max.P1A – Pectoral anter�or marg�n 13.6 12.8 15.5 15.0 11.7 16.1P1I – Pectoral �nner marg�n 8.9 6.8 8.8 8.2 7.0 7.5P1B – Pectoral base length 5.0 5.1 6.2 6.8 5.0 6.1P1P – Pectoral poster�or marg�n 9.6 8.5 11.8 11.0 7.6 11.4P2L – Pelv�c length 10.1 8.6 11.0 10.8 9.6 10.3P2H – Pelv�c he�ght 3.3 3.6 4.8 5.6 4.0 4.9P2I – Pelv�c �nner marg�n 3.7 3.6 5.9 5.8 2.0 3.1CDM – Dorsal caudal marg�n 22.6 19.8 23.3 22.6 21.2 21.3CPV – Preventral caudal marg�n 12.2 10.9 13.2 12.3 10.2 12.2CPU – Upper postventral caudal marg�n 15.9 14.5 17.8 16.4 13.2 16.2CPL – Lower postventral caudal marg�n 3.6 3.9 5.9 4.8 3.4 5.6CFW – Caudal fork w�dth 6.7 6.4 7.7 6.7 5.9 6.7CFL – Caudal fork length 10.8 9.0 10.8 9.2 9.3 10.3HANW – Head w�dth at nostr�ls 7.7 7.0 8.7 7.7 7.6 7.7HAMW – Head w�dth at mouth 10.6 9.7 11.9 11.5 10.1 10.8HDW – Head w�dth 12.1 11.3 13.9 14.8 11.5 13.8TRW – Trunk w�dth 10.9 9.3 12.5 – 8.2 10.7ABW – Abdomen w�dth 8.2 8.1 13.2 – 6.4 9.6TAW – Ta�l w�dth 5.3 5.6 7.0 6.3 4.7 6.7CPW – Caudal peduncle w�dth 2.7 2.6 3.5 2.5 2.4 3.1HDH – Head he�ght 9.1 9.0 12.1 8.5 7.5 11.7TRH – Trunk he�ght 12.0 8.9 13.4 – 7.9 9.1ABH – Abdomen he�ght 12.6 9.3 13.0 – 7.7 8.4TAH – Ta�l he�ght 6.6 6.0 7.3 7.2 5.3 6.2CPH – Caudal peduncle he�ght 2.5 2.2 2.8 2.6 2.3 2.5CLO – Clasper outer length 1.4 4.5 5.6 – 1.7 2.6CLI – Clasper �nner length 5.4 7.8 8.8 – 5.2 6.0CLB – Clasper base w�dth 0.8 1.4 1.9 – 0.9 1.1

Morphometr�c and mer�st�c data were taken from 9 spec�mens of S. montalbani (CSIRO H 2575–02, CSIRO H 2606–02, CSIRO H 2606–05, CSIRO H 5857–06, CSIRO H 5875–07, CSIRO H 5888–03, CSIRO H 5889–20, SUML F 1198 and SUML unreg BRU 136); morphometr�cs only from another 6 spec�mens (USNM 70256 (holotype), CSIRO H 2605–04, CSIRO H 4623–04, CSIRO H 4623–05, QM I 38075 and AMS I 43982–001); and mer�st�cs only from another 10 spec�mens (CSIRO H 1203–02, CSIRO H 1290–02, CSIRO H 1348–01, CSIRO H 2606–06, CSIRO H 4623–02, CSIRO 4708–01, CSIRO H 5889–10, CSIRO H 5889–19, QM I 21518 and WAM P 32843–001). Morphometr�c and mer�st�c deta�ls were taken from the holotype (CSIRO H 4775–01) and two paratypes of the new spec�es (CSIRO H 1405–01, CSIRO H 1662–01); morphometr�cs only from 7 other paratypes (CSIRO H 2867–02, CSIRO

H 2867–03, CSIRO H 2867–04, CSIRO H 2867–05, CSIRO H 2966–01, CSIRO H 5941–01 and NMV A 29563–001); and mer�st�cs from 5 other paratypes (CSIRO CA 121, CSIRO H 1350–02 (4 embryos)).

Spec�mens exam�ned are depos�ted �n the Austral�an Nat�onal F�sh Collect�on, Hobart (CSIRO), the Austral�an Museum, Sydney (AMS), the Nat�onal Museum of V�ctor�a (NMV), the Queensland Museum (QM), the Western Austral�an Museum (WAM), the Museum Zoolog�cum Bogor�ense, Jakarta (MZB), the Hokka�do Un�vers�ty, Faculty of F�sher�es, Hakodate (HUMZ), the Cal�forn�a Academy of Sc�ences, San Franc�sco (CAS, �nc. SU) and the Nat�onal Museum of Natural H�story, Wash�ngton, D.C. (USNM); the�r reg�strat�on numbers are prefixed with these acronyms.

58

Figure 1. Lateral v�ew of: A. Squalus montalbani from NW Austral�a (CSIRO H 2575–02, adult male 681 mm TL); B. Squalus montalbani from Indones�a (MZB 15424, female 945 mm TL); C. Squalus philippinus (=montalbani) holotype (USNM 70256, �mmature male 311 mm TL); d. Squalus mitsukurii holotype (SU 12793, female 719 mm TL).

A

B

C

D

Squalus montalbani Wh�tley, 1931

F�gs 1a–c, 2a,b, 3–5a; Table 1, 2 Squalus philippinus Sm�th and Radcl�ffe, 1912 (jun�or homonym of S. philippinus Shaw, 1804 = Heterodontus portusjacksoni (Meyer, 1793)). In: Smith 1912: 677, pl. 51, fig. 1, Proc. U. S. Natl. Mus. v. 41 (no. 1877). Holotype (un�que): USNM 70256. Squalus montalbani Wh�tley 1931:310. In: Aust. Zool. v. 6 (pt 4). Replacement name for Squalus philippinus Sm�th and Radcl�ffe 1912.

Squalus sp. 1: Wh�te et al., 2006, Economically Important Sharks and Rays of Indonesia, pp 68–69.

Holotype. USNM 70256, �mmature male 311 mm TL, off Sombrero I., west coast of Luzon Island, 13°45′ N, 120°46′ E, Philippines, Albatross sta. 5111, ca. 425 m.Other material. 32 spec�mens. CSIRO H 1290–02, �mmature male 592 mm TL, east of Fl�nders Reef, Queensland, 17°38′ S, 149°23′ E, 600 m; CSIRO H 1348–01, female 375 mm TL, north-west of Saumarez Reef, Queensland, 21°20′ S, 153°32′ E, 502 m; QM


I 21518, female 370 mm TL, east of Capr�corn Group, Queensland, 23°21′ S, 153°56′ E, 460 m; AMS I 20301–027, 310 mm TL, east of Wool�, New South Wales, 29°53′ S, 153°42′ E, 502 m; CSIRO H 4623–02, female 485 mm TL, CSIRO H 4623–04, adult male 811 mm TL, CSIRO H 4623–05, female 760 mm TL, east of Terr�gal, New South Wales, 33°28′ S, 152°04′ E, 383 m; CSIRO H 2606–02, female 557 mm TL, CSIRO H 2606–05, female 607 mm TL, CSIRO H 2606–06, adolescent male 440 mm TL, west of Rottnest Island, Western Austral�a, 32°02′ S, 114°54′ E, 670 m; CSIRO H 2605–04, female 825 mm TL, north-west of Rottnest Island, Western Australia, 31°44′ S, 114°59′ E, 485 m; WAM P 32843–001, �mmature male 440 mm TL, west of Greenhead, Western Australia, 29°59′ S, 114°26′ E, 490 m; CSIRO H 2574–04, �mmature female 221 mm TL, west of Freycinet Estuary, Western Australia, 26°35′ S, 112° 29′ E, 508 m; CSIRO H 2575–02, adult male 681 mm TL, west of Freyc�net Estuary, Western Austral�a, 26° 40′ S, 112°32′ E, 478 m; CSIRO H 1203–02, embryo 208 mm TL, north of Sahul Banks, T�mor Sea, Western Australia, 11°33′ S, 124°58′ E, 415 m; MZB 15018, female 862 mm TL, MZB 15019, adult male 862 mm TL, Cilacap fish landing site, Central Java, Indonesia, 07° 40′ S, 109°00′ E; CSIRO H 5857–06, adult male 678 mm TL, CSIRO H 5889–20, adult male 627 mm TL, CSIRO H 5888–03, female 801 mm TL, CSIRO H 5889–10, female 495 mm TL, CSIRO H 5889–19, female 481 mm TL, AMS I 43982–001, adult male 564 mm TL, QM I 38075, female 801 mm TL, QM I 38076, �mmature male 463 mm TL, MZB 15421, female 912 mm TL, MZB 15424, female 945 mm TL, NMV A 29561–001, female 528 mm TL, Kedonganan fish landing site, Bali, Indonesia, 08°45′ S, 115°10′ E; CSIRO H 5875–07, female 843 mm TL, MZB 15099, female 781 mm TL, Tanjung Luar fish landing site, Lombok, Indonesia, 08°45′ S, 116°35′ E; SUML F 1198, female 694 mm TL, SUML unreg BRU 136, female 520 mm TL, Al�guay Island, Ph�l�pp�nes.

DIAGNOSIS.— A large spec�es of Squalus of the ‘m�tsukur�� group’ w�th the follow�ng comb�nat�on of characters: body elongate to robust, trunk depth 8.9–13.4% TL (mean 11.4% TL, n=14); snout broadly tr�angular, mouth w�dth 1.69–2.32 (1.85) t�mes hor�zontal prenarial length; pre-first dorsal length 26.5–30.7 (29.0)% TL; pre-second dorsal length 57.6–62.8 (60.8)% TL; �nterdorsal space 21.7–25.9 (23.7)% TL; low raked dorsal fins; second dorsal-fin length 11.1–13.9 (12.4)% TL, he�ght 3.4–4.6 (4.0)% TL, �nner marg�n length 4.0–5.9 (4.8)% TL; second dorsal-fin base 15.8–21.3 (20.8) t�mes base of second dorsal sp�ne; prepectoral length 20.8–22.9 (22.0)% TL; pelv�c–caudal space 22.9–26.0 (24.0)% TL; caudal bar almost upr�ght, extend�ng broadly from the caudal fork up the poster�or marg�n of the upper lobe for about 0.6 of �ts length �n �mmatures, upper caudal fr�nge form�ng a deep saddle along m�d-length of lobe; flank denticles tricuspid; 41–47 (mainly 42–44) monospondylous centra, 79–85 precaudal centra, 105–114 total centra; adult s�ze more than 84 cm TL.

A

B

C

Figure 2. Ventral v�ew of the head of: A. Squalus montalbani (CSIRO H 2575–02, adult male 681 mm TL); B. Squalus philippinus (=montalbani) holotype (USNM 70256, �mmature male 311 mm TL); C. Squalus mitsukurii holotype (SU 12793, female 719 mm TL).

60

REMARKS.— Squalus montalbani �s s�m�lar �n appearance to S. mitsukurii and both spec�es occur together �n the same general geograph�c reg�on. Hence, �t �s not unsurpr�s�ng that these forms have been cons�dered to be conspecific (Compagno, 1984). These species differ subtly in caudal fin coloration and these markings are more str�k�ngly obv�ous �n young (F�g. 5) than �n adult spec�mens. The dark caudal bar, wh�ch extends d�agonally as a l�near mark�ng from the or�g�n of the lower lobe to the ax�l of the caudal fork �n S. mitsukurii (exposed along the poster�or caudal marg�n for about half an eye d�ameter), �s less developed than �n S. montalbani (extend�ng broadly from the caudal fork and follow�ng the poster�or marg�n of the upper lobe for 0.6–0.7 of �ts length to an upper level typ�cally demarcated by a posterior projection of the distal part of a fleshy portion of the fin). Also, the dark blotch on the upper caudal lobe �s located more d�stally than �n S. montalbani wh�ch �s represented as a saddle-l�ke extens�on of the upper caudal fr�nge. The late embryo paratypes of S. mitsukurii (SU 7748, 228–237 mm TL) are sl�ghtly larger than the post-natal young of S. montalbani (CSIRO H 1203–02, 207 mm TL) suggest�ng that they are born at d�fferent s�zes. These spec�mens also d�splay sl�ght d�fferences �n the form of the anterior nasal flap and pectoral fin which are either attributable to interspecific or ontogenetic d�fferences. It should be noted that, although paratype SU 7748 cons�sts of 8 late-term embryos labeled as “from uterus of type”, the l�terature suggests that �t may actually conta�n embryos from several �nd�v�duals collected w�th the holotype (Jordan and Snyder, 1903). However, the 5 (of 8) embryos exam�ned �n th�s study all appear to be conspecific based on their morphology and coloration and as such are representat�ve of S. mitsukurii.

These spec�es also d�ffer morphometr�cally (Table 1). The female holotype (SU 12793, 719 mm TL, F�g. 1d, 2c) and �mmature male paratype (SU 7184, 266 mm TL) of Squalus mitsukurii d�ffer from the holotype (USNM 70256, 311 mm TL, F�g. 1c, 2b) and other mater�al of

Figure 3. Cusps of the flank denticles of Squalus montalbani (CSIRO H 2575–02, adult male 681 mm TL). F�eld of v�ew w�dth 1.0 mm.

Figure 4. Lateral view of the dorsal fins of Squalus montalbani (CSIRO H 2575–02, adult male 681 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B

S. montalbani �n the follow�ng rat�os: total length 3.10–3.23 vs. 3.26 (3.33–3.78, n=14) times pre-first dorsal length �n S. montalbani; pre-first dorsal length 1.45–1.73 vs. 1.42 (1.10–1.30) t�mes �nterdorsal space; prepectoral length 1.09–1.28 vs. 1.05 (0.85–1.00) t�mes �nterdorsal space; prepectoral length 1.02–1.07 vs. 0.87 (0.85–0.99) t�mes pelv�c–caudal space; and head he�ght 0.58–0.65 vs. 0.76 (0.66–0.96) t�mes �ts w�dth. Squalus mitsukurii also appears to have sl�ghtly longer head measurements (d�rect head length 23.4–23.5 vs. 23.2 (21.3–23.4)% TL, prepectoral length 23.3–23.9 vs. 22.8 (20.8–22.9)% TL, prenar�al length 5.4–5.5 vs. 4.6 (4.0–5.3)% TL, preoral length 10.6–10.8 vs. 10.1 (8.7–10.6)% TL) and a more slender body (head he�ght 7.5–8.5 vs. 9.1 (9.0–12.1)% TL, trunk he�ght 7.9 vs. 12.0 (8.9–13.4)% TL, abdomen he�ght 7.7 vs. 12.6 (9.3–13.0)% TL, trunk w�dth 8.2 vs. 10.9 (9.3–12.5)% TL, abdomen w�dth 6.4 vs. 8.2 (8.1–13.2)% TL, and caudal peduncle w�dth 2.4–2.5 vs. 2.7 (2.6–3.5)% TL). The mouth of S. mitsukurii appears to be relat�vely narrow compared to S. montalbani (mouth w�dth 6.2–6.3 vs. 6.5 (7.1–8.5)% TL) but th�s may be due to methodolog�cal �ncons�stenc�es. D�fferences between the spec�es also ex�st �n the number of vertebral centra. Chen et al. (1979) prov�ded counts and morphometr�cs for 54 spec�mens of S. mitsukurii from central and northeastern Japan, close to the type local�ty (M�sak�). The�r counts for S. mitsukurii are h�gher (45–51 monospondylous centra, 87–93 precaudal centra, 118–127 total centra) than those


Table 2. Proport�onal d�mens�ons as percentages of total length for S. montalbani from the Ph�l�pp�nes �nclud�ng holotype (USNM 70256), Indones�a and Austral�a.

Philippines Indonesia Australia n = 2 n = 6 n = 6

Holotype Mean M�n. Max. Mean M�n. Max. Mean M�n. Max.TL – Total length 311 607 520 694 719 564 843 707 557 825PCL – Precaudal length 76.5 78.3 78.2 78.3 78.2 77.3 79.9 77.9 77.1 78.9PD2 – Pre-second dorsal length 57.6 60.4 60.2 60.6 61.4 59.8 62.8 60.3 59.1 62.1PD1 – Pre-first dorsal length 30.7 29.8 29.8 29.9 29.0 28.6 30.0 28.6 26.5 29.4SVL – Pre-vent length 47.2 48.8 48.6 49.0 50.4 48.8 52.7 50.3 48.7 51.5PP2 – Prepelv�c length 43.8 48.1 47.7 48.4 48.7 47.2 50.8 48.7 47.5 50.5PP1 – Prepectoral length 22.8 21.5 20.8 22.3 22.1 21.2 22.9 22.1 21.0 22.8HDL – Head length 23.2 22.0 21.7 22.4 22.4 21.7 23.2 22.6 21.3 23.4PG1 – Prebranch�al length 20.3 18.2 17.9 18.4 18.7 17.6 19.3 19.2 18.3 19.7PSP – Presp�racular length 13.4 11.6 11.3 11.8 12.0 11.4 12.5 12.7 11.9 13.6POB – Preorb�tal length 7.1 6.9 6.8 7.1 7.4 6.7 7.7 7.7 7.3 8.1PRN – Prenar�al length 4.6 4.3 4.0 4.5 4.9 4.6 5.1 5.2 4.9 5.3POR – Preoral length 10.1 9.0 8.7 9.4 9.6 9.2 10.2 10.0 9.3 10.6INLF – Inner nostr�l-lab�al furrow space 5.3 4.3 4.2 4.5 4.6 4.5 4.9 4.7 4.5 4.9MOW – Mouth w�dth 6.5 7.7 7.6 7.8 7.5 7.1 7.8 8.0 7.5 8.5ULA – Lab�al furrow length 2.5 2.2 2.2 2.2 2.2 1.9 2.5 2.2 2.1 2.2INW – Internar�al space 5.2 4.4 4.3 4.5 4.3 4.1 4.5 4.6 4.2 5.0INO – Interorb�tal space 8.7 7.8 7.6 8.0 8.2 7.5 9.1 9.0 8.7 9.6EYL – Eye length 4.3 4.0 3.8 4.2 4.9 4.8 5.2 4.2 4.0 4.6EYH – Eye he�ght 1.6 1.9 1.9 1.9 1.7 1.5 1.9 2.0 1.8 2.3SPL – Sp�racle length 1.5 1.3 1.3 1.3 1.7 1.5 1.8 1.7 1.6 1.9GS1 – F�rst g�ll-sl�t he�ght 1.6 1.7 1.7 1.7 2.0 1.8 2.2 2.0 1.8 2.5GS5 – F�fth g�ll-sl�t he�ght 2.0 1.8 1.7 1.8 2.2 2.0 2.4 2.1 1.9 2.3IDS – Interdorsal space 21.7 23.4 22.9 24.0 24.4 22.8 25.9 23.0 22.1 24.1DCS – Dorsal-caudal space 11.2 10.1 9.7 10.4 10.6 9.7 11.2 10.4 9.9 10.6PPS – Pectoral-pelv�c space 18.9 22.4 21.9 22.9 23.0 19.8 25.1 23.0 22.2 25.4PCA – Pelv�c-caudal space 26.3 23.6 23.4 23.7 24.4 23.0 26.0 23.8 22.9 25.3D1L – F�rst dorsal length 14.2 13.5 13.0 14.0 14.2 12.8 15.1 14.9 14.5 15.9D1A – F�rst dorsal anter�or marg�n 12.4 11.3 11.0 11.6 11.6 10.4 12.5 12.2 11.6 12.6D1B – F�rst dorsal base length 8.3 7.9 7.6 8.3 8.9 8.2 9.3 9.3 8.7 9.8D1H – F�rst dorsal he�ght 6.1 6.5 6.5 6.6 6.3 5.7 6.7 6.8 6.4 7.4D1I – F�rst dorsal �nner marg�n 6.2 5.6 5.5 5.8 5.5 4.9 6.2 5.8 5.1 6.8D1P – F�rst dorsal poster�or marg�n 6.7 7.6 7.4 7.9 7.3 6.9 8.0 8.1 7.5 8.8D1ES – F�rst dorsal sp�ne length 2.1 2.7 2.4 3.0 2.9 2.2 4.0 3.0 2.5 3.5D1BS – F�rst dorsal sp�ne base w�dth 0.7 0.6 0.6 0.6 0.6 0.5 0.6 0.6 0.5 0.6D2L – Second dorsal length 13.4 12.3 11.4 13.2 11.8 11.1 12.6 13.0 12.2 13.9D2A – Second dorsal anter�or marg�n 10.2 9.8 9.2 10.3 8.9 8.5 9.4 10.0 9.0 10.8D2B – Second dorsal base length 8.1 7.5 7.0 7.9 7.2 7.0 7.5 8.0 7.1 8.5D2H – Second dorsal he�ght 4.2 4.0 4.0 4.1 3.9 3.4 4.3 4.1 3.5 4.6D2I – Second dorsal �nner marg�n 5.9 4.8 4.3 5.3 4.5 4.0 5.1 5.1 4.7 5.7D2P – Second dorsal poster�or marg�n 5.2 5.1 5.0 5.1 5.3 4.5 6.0 5.7 5.0 6.3D2ES – Second dorsal sp�ne length 3.6 2.9 2.7 3.2 2.9 2.0 3.8 3.3 2.6 3.9D2BS – Second dorsal sp�ne base w�dth 0.8 0.6 0.6 0.6 0.6 0.5 0.6 0.6 0.5 0.6

62

Table 1. cont’d.

Philippines Indonesia Australia n = 2 n = 6 n = 6

Holotype Mean M�n. Max. Mean M�n. Max. Mean M�n. Max.P1A – Pectoral anter�or marg�n 13.6 14.2 14.0 14.3 13.9 12.8 15.2 14.5 13.7 15.5P1I – Pectoral �nner marg�n 8.9 7.8 7.4 8.1 7.3 6.8 7.7 8.2 7.7 8.8P1B – Pectoral base length 5.0 5.7 5.7 5.7 5.4 5.1 5.8 5.9 5.5 6.2P1P – Pectoral poster�or marg�n 9.6 10.8 10.8 10.8 10.0 8.5 11.4 11.0 9.1 11.8P2L – Pelv�c length 10.1 8.8 8.6 9.0 9.9 9.2 10.4 10.2 9.7 11.0P2H – Pelv�c he�ght 3.3 4.7 4.6 4.8 4.1 3.6 4.5 4.5 4.3 4.8P2I – Pelv�c �nner marg�n 3.7 3.9 3.6 4.3 4.9 3.9 5.9 4.9 4.4 5.6CDM – Dorsal caudal marg�n 22.6 21.8 21.6 22.0 21.0 19.8 21.8 22.2 20.9 23.3CPV – Preventral caudal marg�n 12.2 12.2 11.9 12.5 11.6 10.9 12.1 12.6 11.5 13.2CPU – Upper postventral caudal marg�n 15.9 15.9 15.8 16.1 15.4 14.5 16.4 16.8 15.6 17.8CPL – Lower postventral caudal marg�n 3.6 4.9 4.9 4.9 4.8 3.9 5.9 5.0 4.4 5.5CFW – Caudal fork w�dth 6.7 6.8 6.8 6.8 6.7 6.4 7.0 7.3 7.0 7.7CFL – Caudal fork length 10.8 9.9 9.4 10.4 9.5 9.0 9.8 10.1 9.4 10.8HANW – Head w�dth at nostr�ls 7.7 7.5 7.1 7.8 7.2 7.0 7.4 8.1 7.5 8.7HAMW – Head w�dth at mouth 10.6 10.4 10.0 10.8 10.4 9.7 11.4 11.4 10.6 11.9HDW – Head w�dth 12.1 13.7 13.5 13.9 12.2 11.3 13.0 12.7 11.8 13.2TRW – Trunk w�dth 10.9 10.6 10.5 10.7 10.5 9.3 11.7 11.0 9.9 12.5ABW – Abdomen w�dth 8.2 8.9 8.9 9.0 10.3 8.1 12.6 10.7 8.9 13.2TAW – Ta�l w�dth 5.3 6.0 5.9 6.2 6.4 5.6 7.0 6.5 5.7 7.0CPW – Caudal peduncle w�dth 2.7 2.7 2.6 2.9 3.2 3.1 3.4 3.3 3.2 3.5HDH – Head he�ght 9.1 9.3 9.2 9.4 10.8 9.2 12.0 10.6 9.0 12.1TRH – Trunk he�ght 12.0 9.9 9.2 10.7 11.6 10.1 13.1 11.6 8.9 13.4ABH – Abdomen he�ght 12.6 10.6 10.2 10.9 11.3 9.3 13.0 11.8 9.5 12.9TAH – Ta�l he�ght 6.6 6.5 6.4 6.7 6.6 6.0 7.3 6.8 6.2 7.2CPH – Caudal peduncle he�ght 2.5 2.5 2.5 2.6 2.4 2.2 2.6 2.7 2.6 2.8CLO – Clasper outer length 1.4 – – – 4.9 4.4 5.3 5.0 4.5 5.6CLI – Clasper �nner length 5.4 – – – 7.6 7.4 7.9 8.3 7.8 8.8CLB – Clasper base w�dth 0.8 – – – 1.6 1.4 1.9 1.5 1.4 1.7

of S. montalbani (41–47 monospondylous centra, 79–85 precaudal centra, 105–114 total centra, n=30).

Some m�nor d�fferences ex�st between populat�ons of S. montalbani across the Indo–West Pacific. Vertebral counts var�ed sl�ghtly between mater�al from Indones�a (41–45 (mean 42.8) monospondylous centra, 79–83 (mean 80.5) precaudal centra, 105–111 (mean 107.6) total centra, n=12) and northern Austral�a (42–47 (mean 43.6) monospondylous centra, 81–85 (mean 83.1) precaudal centra, 110–114 (mean 112.2) total centra, n=14). Morphometr�cs from the three populat�ons are reasonably cons�stent (Table 2) w�th m�nor var�at�on �n eye length, first dorsal-fin base length, pelvic-fin inner marg�n length, sp�racle length, caudal peduncle w�dth, and head w�dth at the anter�or nostr�ls. Indones�an mater�al �s darker �n overall colorat�on than Austral�an

mater�al but the general morphology of spec�mens from both reg�ons and the Ph�l�pp�nes �s s�m�lar.

Squalus chloroculus sp. nov.


Squalus mitsukurii: (not Jordan and Snyder) Last and Stevens, 1994, Sharks and Rays of Australia, pp 49, 101, figs 8.24, 8.39, pl. 5; Gomon, Glover and Ku�ter, 1994, The Fishes of Australia’s South Coast, pp 105, 107, 108, figs 56, 57.Holotype. CSIRO H 4775–01, adult male 753 mm TL, off Portland, V�ctor�a, 38° S, 141° E, 17 Apr�l 1998.Paratypes. 14 spec�mens. CSIRO CA 121, female 731 mm TL, off Ulladulla, New South Wales, 35°27′ S, 150°51′ E, 400 m; NMV A 29563–001, adult male


A

B

Figure 5. Juvenile coloration of the caudal fin of: A. Squalus montalbani (CSIRO H 1203–02, �mmature female 207 mm TL); B. Squalus mitsukurii paratype (CAS SU 7748, female embryo 237 mm TL).

856 mm TL, off St Helens, Tasman�a, ca. 42° S, 148° E, 550 m; CSIRO H 5941–01, adult male 762 mm TL, west of Cape Sorell, Tasmania, 42°10′ S, 144°45′ E, 460 m; CSIRO H 1662–01, adult male 722 mm TL, east of Mar�a Island, Tasmania, 42°40′ S, 148°24′ E, 468 m; CSIRO H 1350–02, 4 embryos 217–238 mm TL, northwest of Macquarie Harbour, Tasmania, 41°52′ S, 144°23′ E, 1370 m; CSIRO H 1405–01, adult male 760 mm TL, south of King Island, Tasmania, 40°46′ S, 143°42′ E, 216 m; CSIRO H 2966–01, female 832 mm TL, Great Australian Bight, South Australia, 33°47′ S, 131° 27′ E, 780 m; CSIRO H 2867–02, adult male 733 mm TL, CSIRO H 2867–03, adult male 685 mm TL, CSIRO H 2867–04, female 797 mm TL, CSIRO H 2867–05, female 789 mm TL, Great Austral�an B�ght, South Australia, 33°25′ S, 129°54′ E, 514 m.

DIAGNOSIS.— A large spec�es of Squalus of the ‘m�tsukur�� group’ w�th the follow�ng comb�nat�on of characters: body moderately robust, trunk depth 10.4–13.8% TL (mean 11.7% TL, n=9); snout broadly tr�angular, mouth w�dth 1.72–2.48 (2.07) t�mes hor�zontal prenar�al length; pre-first dorsal length 29.2–31.8 (30.2)% TL; pre-second dorsal length 60.7–63.6 (61.7)% TL; �nterdorsal

space 23.7–27.5 (24.9)% TL; low raked dorsal fins; second dorsal-fin length 10.9–12.2 (11.6)% TL, height 3.4–4.0 (3.7)% TL, �nner marg�n length 3.9–5.0 (4.4)% TL; second dorsal-fin base 15.1–20.0 (17.2) times base of second dorsal sp�ne; prepectoral length 21.1–24.3 (22.2)% TL; pelv�c–caudal space 21.8–25.0 (23.5)% TL; caudal bar almost upr�ght, extend�ng broadly from the caudal fork up the poster�or marg�n of the upper lobe for 0.6–0.7 of �ts length �n �mmatures, upper caudal fr�nge forming a narrow saddle along mid-length of lobe; flank dent�cles tr�cusp�d; 43–46 monospondylous centra, 84–86 precaudal centra, 111–115 total centra; adult max�mum s�ze at least 85 cm TL.

DESCRIPTION.— Body fus�form, moderately elongate to robust, nape prominently humped; deepest near first dorsal-fin spine, maximum depth 0.89 (0.88–1.13 in paratypes) t�mes w�dth; trunk depth 1.03 (0.98–1.11) t�mes abdomen depth; head moderately elongate, 23.0 (22.1–23.8)% TL; caudal peduncle robust, 24.0 (21.8–25.0)% TL. Head robust, rather broad, w�dth 1.12 (1.07–1.32) t�mes trunk w�dth, 1.30 (1.05–1.37) t�mes abdomen w�dth, sl�ghtly depressed forward of sp�racles, becom�ng subtriangular in cross-section towards pectoral-fin origin, length 2.21 (2.15–2.33) �n pre-vent length; he�ght 0.77 (0.67–0.85) t�mes w�dth. Snout moderately elongate, narrowly tr�angular �n lateral v�ew, apex bluntly po�nted, lateral prenar�al marg�n weakly angular; narrowly po�nted �n dorsal v�ew, hor�zontal length 1.64 (1.61–1.62) t�mes eye length, 0.90 (0.76–0.81) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.20 (2.16–2.23) t�mes �n preoral length. Eye oval, of moderate s�ze, length 5.25 (4.98–5.37) �n head, 2.13 (1.90–2.20) t�mes he�ght; strongly notched poster�orly, notch deep anter�orly, becom�ng shallow near anteroventral marg�n of sp�racle (not connected to sp�racle �n some paratypes). Sp�racle moderate, broadly crescent�c but var�able; broad, lobe-l�ke fold on poster�or marg�n; greatest d�ameter 2.68 (2.22–3.69) �n eye length. G�ll open�ngs almost upr�ght, first 4 subequal in size, fifth longest, height of fifth slit 2.1 (1.9–2.5)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.28 (1.09–1.30) �n preoral length; prom�nent postoral groove, much longer than upper lab�al furrows (rarely subequal), extend�ng posterolaterally from angle of jaws; 1 (1–2) ser�es of funct�onal teeth �n upper jaw, 2 (2–3) ser�es �n lower. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Nostr�ls small, almost transverse; anterior nasal flap weakly bilobed or s�ngle-lobed; anter�or lobe large, narrowly rounded, subtriangular (flattened in some paratypes); posterior lobe of nasal flap either indistinguishable, rudimentary or weak �n paratypes (somet�mes var�able w�th�n an �nd�v�dual); �nternar�al space 2.23 (2.93–2.21) �n preoral length, 2.65 (2.48–2.83) t�mes nostr�l length. Dermal dent�cles (based on adult male CSIRO H 1405–01) on flank very small, strongly imbricate, tricuspidate; crown broad w�th pronounced med�an r�dge; med�an r�dge

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Figure 6. Lateral v�ew of Squalus chloroculus sp. nov. holotype (CSIRO H 4775–01, adult male 753 mm TL).

Figure 7. Ventral v�ew of the head of Squalus chloroculus sp. nov. holotype (CSIRO H 4775–01, adult male 753 mm TL).

commenc�ng well anter�or of rest of crown, w�th a mes�al furrow develop�ng anter�orly and converg�ng towards poster�or t�p of crown; d�stal r�dge extend�ng laterally along crown, lateral cusps short or var�ably developed, low, weak, w�th very narrow mes�al furrow. F�rst dorsal fin small, low, strongly raked, very broadly rounded ap�cally; anter�or marg�n strongly convex; poster�or marg�n weakly concave, upper port�on subvert�cal (usually d�rected sl�ghtly or strongly anterodorsally from bottom to top �n paratypes); lower half almost stra�ght or weakly concave; free rear t�p relat�vely th�ck basally, very short; inner margin of fin almost straight; insertion of base extremely well forward of pelvic-fin origin, well posterior to free rear tip of pectoral fin; fin-spine origin slightly posterior to pectoral-fin insertion (more

poster�or �n some paratypes); sp�ne base moderately broad, 0.7 (0.6–0.8)% TL, exposed anter�orly well below junction of spine and soft portion of fin; soft portion of fin connected well above mid-point of total spine length (sp�ne often damaged); sp�ne taper�ng d�stally (often abraded ap�cally), anter�or marg�n almost stra�ght; exposed port�on only sl�ghtly raked, much shorter �n length than exposed portion of second dorsal-fin spine (also marginally abraded apically); pre-first dorsal length 3.43 (3.15–3.42) times in TL; first dorsal-fin length 2.41 (2.14–2.41) t�mes �ts he�ght, 1.25 (1.17–1.26) t�mes second dorsal-fin length; first dorsal-fin height 1.50 (1.59–1.78) times second dorsal-fin height; exposed first dorsal spine length 0.56 (0.39–0.50) times height of fin. Second dorsal fin small, strongly raked; anterior margin


Table 3. Proport�onal d�mens�ons as percentages of total length for the holotype (CSIRO H 4475–01) and ranges for the 9 paratypes of Squalus chloroculus sp. nov.

S. chloroculus sp. nov. Paratypes

Holotype M�n. Max. Mean

TL – Total length 753 721 856 783PCL – Precaudal length 78.8 77.9 79.7 78.6PD2 – Pre-second dorsal length 61.8 60.7 63.6 61.7PD1 – Pre-first dorsal length 29.2 29.3 31.8 30.2SVL – Pre-vent length 50.7 49.5 52.1 50.8PP2 – Prepelv�c length 49.0 47.9 51.0 49.3PP1 – Prepectoral length 22.3 21.1 24.3 22.2HDL – Head length 23.0 22.0 23.8 22.7PG1 – Prebranch�al length 19.4 18.5 20.1 19.2PSP – Presp�racular length 13.0 12.3 13.6 12.8POB – Preorb�tal length 7.5 7.5 8.1 7.7PRN – Prenar�al length 5.3 4.7 5.3 5.0POR – Preoral length 9.9 9.5 10.1 9.8INLF – Inner nostr�l-lab�al furrow space 5.0 4.4 4.9 4.7MOW – Mouth w�dth 7.7 7.7 9.1 8.1ULA – Lab�al furrow length 2.4 2.0 2.5 2.3INW – Internar�al space 4.5 4.5 5.1 4.8INO – Interorb�tal space 8.0 8.3 9.5 8.8EYL – Eye length 4.4 4.3 4.5 4.4EYH – Eye he�ght 2.1 2.0 2.3 2.2SPL – Sp�racle length 1.6 1.2 2.0 1.5GS1 – F�rst g�ll-sl�t he�ght 1.9 1.7 2.8 2.1GS5 – F�fth g�ll-sl�t he�ght 2.1 1.9 2.5 2.2IDS – Interdorsal space 25.2 23.7 27.5 24.9DCS – Dorsal-caudal space 10.2 9.3 10.7 10.1PPS – Pectoral-pelv�c space 24.0 21.6 25.2 23.2PCA – Pelv�c-caudal space 24.0 21.8 25.0 23.5D1L – F�rst dorsal length 14.5 13.1 14.8 14.2D1A – F�rst dorsal anter�or marg�n 12.2 10.6 12.5 11.5D1B – F�rst dorsal base length 9.3 8.1 9.6 8.7D1H – F�rst dorsal he�ght 6.0 5.9 6.8 6.2D1I – F�rst dorsal �nner marg�n 5.3 5.1 6.1 5.4D1P – F�rst dorsal poster�or marg�n 6.7 6.8 8.2 7.4D1ES – F�rst dorsal sp�ne length 3.4 2.3 3.3 2.9D1BS – F�rst dorsal sp�ne base w�dth 0.7 0.6 0.8 0.7D2L – Second dorsal length 11.6 10.9 12.2 11.6D2A – Second dorsal anter�or marg�n 8.9 8.0 9.6 8.9D2B – Second dorsal base length 7.4 6.5 7.4 7.2D2H – Second dorsal he�ght 4.0 3.4 3.9 3.7D2I – Second dorsal �nner marg�n 4.5 3.9 5.0 4.4D2P – Second dorsal poster�or marg�n 5.1 4.8 6.1 5.2D2ES – Second dorsal sp�ne length 4.0 2.5 3.9 3.2D2BS – Second dorsal sp�ne base w�dth 0.7 0.6 0.8 0.7

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Table 1. cont’d.

S. chloroculus sp. nov. Paratypes

Holotype M�n. Max. MeanP1A – Pectoral anter�or marg�n 14.9 13.7 16.9 14.9P1I – Pectoral �nner marg�n 7.6 7.0 8.6 7.8P1B – Pectoral base length 5.4 5.1 6.0 5.6P1P – Pectoral poster�or marg�n 11.0 10.2 12.8 11.2P2L – Pelv�c length 10.5 9.2 10.5 9.8P2H – Pelv�c he�ght 5.1 4.4 5.4 5.0P2I – Pelv�c �nner marg�n 5.3 1.9 4.6 3.3CDM – Dorsal caudal marg�n 20.9 19.2 21.7 21.0CPV – Preventral caudal marg�n 12.2 11.0 12.8 11.7CPU – Upper postventral caudal marg�n 15.8 14.7 16.5 15.8CPL – Lower postventral caudal marg�n 5.1 5.2 6.3 5.5CFW – Caudal fork w�dth 7.1 6.6 7.2 7.0CFL – Caudal fork length 9.5 9.1 9.7 9.4HANW – Head w�dth at nostr�ls 7.9 7.6 8.5 7.9HAMW – Head w�dth at mouth 11.1 10.8 12.5 11.6HDW – Head w�dth 13.3 12.7 14.9 13.7TRW – Trunk w�dth 11.8 10.5 13.0 11.7ABW – Abdomen w�dth 10.2 10.1 13.2 11.5TAW – Ta�l w�dth 6.1 4.8 7.3 6.3CPW – Caudal peduncle w�dth 3.3 3.0 3.7 3.2HDH – Head he�ght 10.2 9.4 11.5 10.3TRH – Trunk he�ght 10.5 10.4 13.8 11.7ABH – Abdomen he�ght 10.3 10.1 13.5 11.5TAH – Ta�l he�ght 6.4 6.0 6.8 6.4CPH – Caudal peduncle he�ght 2.7 2.3 2.6 2.5CLO – Clasper outer length 3.8 4.1 4.6 4.3CLI – Clasper �nner length 7.2 6.9 8.8 7.7CLB – Clasper base w�dth 1.5 1.5 1.7 1.6

convex, apex broadly rounded; poster�or marg�n strongly concave, max�mum concav�ty just above m�d-po�nt of marg�n (var�able �n paratypes), upper port�on almost vert�cal (d�rected sl�ghtly dorsoposter�orly �n paratypes CSIRO H 2867–05 and H 2966–01); free rear t�p very elongate, inner margin length 1.12 (1.06–1.39) times fin height; second dorsal-fin length 2.89 (2.97–3.44) times its height; spine length 1.01 (0.68–1.06) in height of fin; fin-spine origin well behind free rear tip of pelvic fin, exposed near level of junction with spine and soft portion of fin (connected sl�ghtly above �n some paratypes); second sp�ne moderately broad based, 0.7 (0.6–0.8)% TL, sharply po�nted d�stally when undamaged, usually abraded; sp�ne t�p when undamaged extend�ng to about level of insertion of fin; soft portion and spine apices subequal �n he�ght; �nterdorsal space 0.89 (0.81–0.99) �n length

from snout tip to pectoral-fin origin, 1.16 (1.10–1.30) in pre-first dorsal length; interdorsal ridge rudimentary or absent, more obvious posteriorly. Pectoral fin large, more so �n large females (anter�or marg�n 15.7–16.9% TL, n=3) than males (anter�or marg�n 13.7–14.9% TL, n=7 �nc. holotype), anter�or marg�n weakly convex; �nner marg�n moderately convex, length 7.6 (7.0–8.6)% TL; apex narrowly rounded (var�able �n some male paratypes, significantly broader), weakly lobe-like and not falcate; poster�or marg�n weakly to moderately concave; free rear t�p narrowly rounded to weakly angular; base very short, 2.76 (2.35–3.08) �n length of anter�or marg�n. Pelv�c fins large, subtriangular, anterior and posterior margins almost stra�ght, apex broadly rounded, free rear t�p acute. Clasper extend�ng well beyond free rear t�p of pelv�c fin. Tail long, subcircular in cross-section anteriorly,


tapering slightly to second dorsal fin, beyond second dorsal fin tapering more rapidly, moderately depressed, broadly sem�c�rcular poster�orly; ventral groove well developed, broad, shallow, w�th obv�ous med�al r�dge (better developed poster�orly); lateral keels very well developed, or�g�nat�ng under (or sl�ghtly poster�or to) insertion of second dorsal fin, terminating about half an eye diameter behind caudal-fin insertion; pelvic–caudal space 1.00 (0.94–1.11) �n pectoral–pelv�c space, 0.93 (0.87–1.11) �n prepectoral length; dorsal–caudal space 2.46 (2.31–2.65) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal p�t moderate to weak. Caudal fin relatively long; upper lobe relatively broad, t�p narrowly rounded, postventral marg�n weakly convex (much more pronounced �n CSIRO H 5941–01); lower lobe acute (more broadly rounded �n CSIRO H 1662–01); dorsal caudal marg�n 1.10 (1.03–1.16) �n head length; length of lower caudal lobe 1.71 (1.66–1.89) �n upper lobe length. Vertebral centra 114 (111–115 �n 7 paratypes), monospondylous 43 (43–46), precaudal 84 (84–86) and caudal 30 (27–30). Teeth �n upper jaw (�n paratype CSIRO H 1662–01) 14+15=29, lower jaw 12+12=24.

COLOUR.— When fresh (based on holotype): rather un�formly grey�sh; undersurface of head paler grey, l�ght and dark tonal areas sharply demarcated, significantly paler ventrally from snout to outer edge of sp�racle, along subocular r�dge to m�d-g�ll sl�ts; rest of belly paler than dorsal surface, almost white. Dorsal fins mostly pale grey, distinct blackish margin extending from above fin spine and along outer marg�n to notch �n poster�or marg�n (some var�at�on as dark marg�n not obv�ous �n some paratypes). Second dorsal fin similar with variable defined blackish extremity; no obvious pale areas at anterior base of fins; fin spines pale, greyish brown to opaque white. Caudal fin mostly greyish; broad dark area at notch, extending dorsally for sl�ghtly longer than an eye d�ameter along upper postdorsal marg�n (usually to a pos�t�on demarcated by a l�ne through the ta�l vertebrae where �t meets the

Figure 8. Cusps of the flank denticles of Squalus chloroculus sp. nov. paratype (CSIRO H 1405–01, adult male 760 mm TL). F�eld of v�ew w�dth 1.3 mm.

upper poster�or caudal marg�n); t�p of upper lobe and upper poster�or marg�n pale; outer marg�n of lower lobe pale. Pectoral fin upper surface greyish with narrow pale poster�or marg�n and apex; ventral surface paler, w�th less well demarcated outer margin. Pelvic fin upper surface similar to upper surface of pectoral fin; posterior margin strongly demarcated pale; ventral surface un�formly pale. Claspers mostly pale, basal dorsal half of clasper grey�sh. Eye green�sh �n l�fe, otherw�se blu�sh black. In preservat�ve (based on holotype): s�m�lar, more un�formly med�um grey above; ventral surface somewhat mottled, sl�ghtly paler grey; undersurface of snout off-wh�te, to mottled grey. Pectoral and pelvic fins greyish dorsally w�th sl�ghtly paler poster�or marg�ns. Late embryo paratypes from Tasman�a (CSIRO H 1350–02, F�g. 10) w�th more strongly demarcated pale and dark tonal areas on head; apical two-thirds of dorsal fins black, anterior basal half of fins and free rear tip white; skin on anterior dorsal-fin spines blackish. Caudal fin pattern strong, black and wh�te; broad black caudal bar over caudal fork, extend�ng up the dorsal lobe for more than half �ts length (as a posterior projection of distal part of fleshy portion of fin), then extending slightly above fleshy portion to jo�n large, d�ffuse-edged, black blotch on central anter�or port�on of upper lobe; large black blotch at base of lower lobe narrowly connected to caudal bar below fleshy portion of fin; apical third of upper lobe and most of ventral lobe v�v�d wh�te.

A

B

Figure 9. Lateral view of the dorsal fins of Squalus chloroculus sp. nov. paratype (CSIRO H 5941–01, adult male 762 mm TL) – A. first dorsal fin, B. second dorsal fin.

68

SIZE.— Females and males reach at least 832 (paratype) and 856 mm TL (paratype), respect�vely; males mature by 685 mm TL.

DISTRIBUTION.— Upper to m�d cont�nental slope off southern Austral�a from New South Wales (ca. 35° S) to the Great Austral�an B�ght (33° S, 129° E). Known from depths of 216–1360 m.

ETYMOLOGY.— Der�ved from the Greek choloros mean�ng ‘green’ and the Lat�n oculus mean�ng ‘eye’ �n allus�on to the v�v�d green eyes ev�dent on th�s spec�es when fresh.

VERNACULAR.— Greeneye Spurdog.

REMARKS.— Squalus chloroculus d�ffers from S. montalbani �n the structure of the CO1 gene (see Ward et al., 2007, Part 12) but the two spec�es are very s�m�lar morpholog�cally. M�nor d�fferences �n colorat�on were detected �n Austral�an ‘m�tsukur��’-l�ke spurdogs dur�ng �nvest�gat�ons for the Austral�an shark and ray gu�de (Last and Stevens, 1994) but these forms were prov�s�onally grouped under S. mitsukurii. Three shape characters d�st�ngu�sh Austral�an populat�ons of the two species: pre-first dorsal length 2.41–2.77 vs. 1.91–2.41 times second dorsal-fin length in S. montalbani; hor�zontal pre-second dorsal length 5.03–5.81 vs. 4.20–4.92 times second dorsal-fin length; and second dorsal-fin length 10.9–12.2 vs. 12.2–13.9% TL. In addition, S. chloroculus has smaller dorsal fins (the mean values for all 6 measurements of both dorsal fins are higher for S. montalbani) with broader fin spine bases (means 0.68 vs. 0.58% TL for first dorsal-fin spine, 0.68 vs. 0.60% TL for second dorsal-fin spine), shorter adult claspers (outer length 3.8–4.6, mean 4.3% TL vs. 4.5–5.6, mean 5.0%

Figure 10. Juvenile coloration of the caudal fin of Squalus chloroculus sp. nov. paratype (CSIRO H 1350–02, �mmature male 225 mm TL).

TL), and the upper postventral caudal marg�n �s short relat�ve to the lower postventral marg�n (rat�o 2.54–3.15, mean 2.88 vs. 2.92–3.89, mean 3.36). Vertebral counts are very s�m�lar, w�th S. chloroculus hav�ng a marg�nally h�gher average precaudal count (84–86, mean 84.8, n=8 vs. 81–85, mean 83.1, n=14). Squalus chloroculus d�ffers from S. mitsukurii in caudal fin coloration (caudal bar upr�ght and marg�nal rather than obl�que) and by hav�ng sl�ghtly lower vertebral counts (43–46 vs. 45–51 monospondylous centra, 84–86 vs. 87–93 precaudal centra, 111–115 vs. 118–127 total centra).

Squalus chloroculus displays some intraspecific var�ab�l�ty �n shape. A large adult male paratype from off St Helens, Tasman�a (NMV A 29563–001), has a much broader upper caudal lobe, more broadly rounded lower caudal lobe, more robust head, and �s more un�formly dark ventrally and dorsally than other types. Its pectoral fin is also less falcate w�th a more broadly rounded apex.

Other material.Squalus mitsukurii: SU 7184 (paratype), �mmature male 266 mm TL, SU 7748 (paratypes), 5 of 8 embryos exam�ned 228–237 mm TL, SU 12793 (holotype), 719 mm TL, M�sak�, Honshu Island, Japan; HUMZ 79797, female 855 mm TL, HUMZ 79798, female 854 mm TL, Kyushu–Palau R�dge, Japan, 320–640 m; HUMZ 101719, adult male 657 mm TL, northwest of Ok�nawa, Japan.

ACKNOWLEDGEMENTS

Th�s study was conducted as part of a project on southern Australian dogfishes funded by the Fisheries Research and Development Trust (FRDC 1998/108). We thank curators and curator�al staff for ass�stance w�th obta�n�ng spec�mens and collect�on data: Mart�n Gomon and D� Bray (NMV), Barry Hutch�ns, Sue Morr�son and Glenn Moore (WAM), Jeff Johnson (QM), Mark McGrouther (AMS), Kazu Nakaya and H�sash� Imamura (HUMZ), Dave Catan�a (CAS), Jeff W�ll�ams (USNM), and Alasta�r Graham (CSIRO). Thanks also go to Lou�se Conboy for ed�ted �mages of the types and other mater�al, Bryn Farmer for supply�ng dent�cle �mages, Tony Rees for ass�st�ng w�th preparat�on of SEMs, John Stevens for obta�n�ng tooth counts and ed�t�ng, T�m Founta�n for prepar�ng rad�ographs and obta�n�ng mer�st�c �nformat�on, Dan�el Gledh�ll for ass�st�ng w�th a l�terature summary, John Pogonosk� for ed�t�ng and techn�cal ass�stance, and also to Terry Walker and Ken Graham. We thank all scientific part�c�pants and crew of several research voyages for the�r ass�stance �n collect�ng mater�al of spec�es treated �n th�s paper. Indones�an mater�al was collected as part of a 5-year ACIAR funded project and �mportant contr�butors to th�s project �nclude, Junaed� (Kedonganan F�sh Market, Bali), processors and fishers at Tanjung Luar fish market (Lombok), Fahm� (LIPI), Dharmad� (RCCF), Jenny G�les (QDPI&F), Ono Kurnaen Sumadh�harga (LIPI), Subhat


Nurhak�m (RCCF), Ian Potter (Murdoch Un�vers�ty), Cathy D�chmont and Steve Blaber (CSIRO).

REFERENCES

Compagno, L.J.V. (1984) FAO spec�es catalogue. Sharks of the World. An annotated and �llustrated catalogue of shark spec�es known to date. Part 1. Hexanch�formes to Lamn�formes. FAO Fisheries Synopsis, No. 125, v. 4 (part 1), pp 1–250.


Gomon, M.F., Glover, J.C.M. and Ku�ter, R.H. (1994) The Fishes of Australia’s South Coast. State Pr�nt, Adela�de, 992 pp.



Jordan, D.S. and Fowler, H.W. (1903) A rev�ew of the

elasmobranchiate fishes of Japan. Proceedings of the United States National Museum, 26, 593–674.

Meyer, F.A.A. (1793) Systemat�sch-summar�sche Uebers�cht der neuesten zoolog�schen Entdeckungen �n Neuholland und Afr�ka. Le�pz�g, Dyk�schen. Zool. Entdeckungen, 1–178.

Shaw, G. (1804) General zoology or systematic natural history. Vol. 5 (pt 1), 1–25 and (pt 2), 251–463.

Sm�th, H.M. (1912) The squalo�d sharks of the Ph�l�pp�ne Arch�pelago, w�th descr�pt�ons of new genera and spec�es. [Scientific results of the Philippine cruise of the Fisheries steamer “Albatross,” 1907–10., No. 15.]. Proceedings of the United States National Museum, 41(1877), 677–685.

Ward, R.D., Holmes, B.H., Zemlak, T.S. and Sm�th, P.J. (2007) Part 12 — DNA barcod�ng d�scr�m�nates spurdogs of the genus Squalus, p. 117–130. In: Descr�pt�ons of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.


Whitley, G.P. (1931) New names for Australian fishes. Australian Zoologist, 6(4), 310–334.


Part 7 — Two new species of Squalus of the ‘mitsukurii group’ from the Indo–Pacific

W�ll�am T. Wh�te, Peter R. Last and John D. Stevens


ABSTRACT. — Two new spec�es of spurdog, Squalus edmundsi sp. nov. and Squalus grahami sp. nov., are descr�bed based on spec�mens from the upper cont�nental slope off western and eastern Austral�a respect�vely. The new spec�es, one of wh�ch has been confused w�th S. mitsukurii from the western North Pacific, belong to a subgroup of Squalus, the ‘m�tsukur�� group’, whose members all have tr�cusp�d denticles, a moderately elongate snout, a dark bar on or reaching the posterior notch of the caudal fin, and a moderately slender body. The two spec�es are clearly separable from each other and the�r closely related congeners by a comb�nat�on of colorat�on, mer�st�cs and morphometr�cs.

Key words. Squalo�dea – Squal�dae – Squalus edmundsi – Squalus grahami – new spec�es – Indo–West Pacific


INTRODUCTION

Last and Stevens (1994) identified 9 Australian Squalus spec�es of wh�ch 6 appear to be formally undescr�bed. Two of these spec�es, referred to as Squalus sp. C and Squalus sp. F (Compagno et al., 2005), belong to the ‘m�tsukur�� group’ but are more slender than most other group members. Recent work on S. sp. C (sensu Last and Stevens, 1994) from Western Austral�a revealed two d�st�nct undescr�bed spec�es: S. altipinnis Last, Wh�te and Stevens, 2007 (Part 5 of th�s �ssue), wh�ch has a h�gh dorsal fin and short snout reminiscent of the ‘megalops group’, and a second spec�es also w�th an upr�ght dorsal fin similar to S. altipinnis, but w�th a longer and more po�nted snout and a ‘m�tsukur��-l�ke’ dark bar on the posterior notch of the caudal fin. Market surveys at various land�ng s�tes �n eastern Indones�a between Apr�l 2001 and March 2006 produced a var�ety of sharks, skates, rays and ch�maeras, �nclud�ng several squal�d spec�es (Wh�te et al., 2006). Amongst th�s mater�al were four spec�es of Squalus, one of wh�ch (�.e. Squalus cf. sp. C, sensu Wh�te et al., 2006) appears to be conspecific with the second undescr�bed, northwestern Austral�an spec�es. Squalus sp. F from eastern Australia, which was identified by research to produce an Austral�an faunal gu�de (Last and Stevens, 1994), rema�ns undescr�bed. These two new members of the ‘m�tsukur�� group’ are descr�bed here�n based on Austral�an mater�al and compar�sons are made w�th other s�m�lar congener�cs.

METHODS

Methods follow those outl�ned �n Part 1 of th�s �ssue (Last et al., 2007). For Squalus edmundsi sp. nov., both morphometr�cs and mer�st�cs were taken from the holotype (CSIRO H 2566–01) and the follow�ng 5 paratypes: CSIRO H 2605–05, CSIRO H 2605–06, CSIRO H 2605–07, CSIRO H 2608–16 and CSIRO H 2590–11 (Table 1). In add�t�on, mer�st�cs were taken from the follow�ng 5 paratypes: CSIRO H 822–16, CSIRO H 1207–06, CSIRO H 2567–10, CSIRO H 2575–18 and CSIRO H 2599–01. For Squalus grahami sp. nov., both morphometr�cs and mer�st�cs were taken from the holotype (CSIRO H 4476–01) and the follow�ng 5 paratypes: CSIRO H 4476–08, CSIRO H 4682–01, CSIRO H 4682–02, CSIRO H 4682–03 and CSIRO H 4682–04 (Table 1). In add�t�on, mer�st�cs were taken from the follow�ng 23 paratypes: CSIRO H 452, CSIRO H 453, CSIRO H 454, CSIRO H 455, CSIRO H 456, CSIRO H 457–01, CSIRO H 602–02, CSIRO H 644–03, CSIRO H 644–05, CSIRO H 644–06, CSIRO H 1311–03, CSIRO H 1312–06, CSIRO H 1312–07, CSIRO H 1312–08, CSIRO H 1312–09, CSIRO H 1344–03, CSIRO H 1346–01, CSIRO H 1347–01, CSIRO H 1406–01, CSIRO H 2468–02, CSIRO H 2469–04, CSIRO H 2469–05 and CSIRO H 2688–02. In each descr�pt�on, morphometr�c and mer�st�c values for the holotype are given first followed in parentheses by the ranges of the paratypes.

Type spec�mens are depos�ted �n the Austral�an Nat�onal

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F�sh Collect�on, Hobart (CSIRO), and at the Austral�an Museum, Sydney (AMS) and Nat�onal Museum of Victoria (NMV); their registration numbers are prefixed w�th these acronyms.

Squalus edmundsi sp. nov.


Squalus sp. C (�n part): Last and Stevens, 1994, Sharks and Rays of Australia, pp 49, 94, fig. 8.33, pl. 6.Squalus sp. C: Compagno, Dando and Fowler, 2005, A Field Guide to the Sharks of the World, pp 79-80, fig, pl 2.Squalus cf. sp. C: Wh�te et al., 2006, Economically Important Sharks and Rays of Indonesia, pp 72–73.

Holotype. CSIRO H 2566–01, adult male 614 mm TL, west of Bernier Island, Western Australia, 24°55′ S, 112°11′ E, 344 m, 28 January 1991.Paratypes. 11 spec�mens. CSIRO H 2608–16, female 732 mm TL, Rottnest Canyon, Western Austral�a, 31°55′ S, 115°10′ E, 850 m; CSIRO H 2605–05, female 649 mm TL, CSIRO H 2605–06, female 560 mm TL, CSIRO H 2605–07, female 508 mm TL, northwest of Rottnest Island, Western Australia, 31°44′ S, 114°58′ E, 485 m; CSIRO H 2599–01, adult male 585 mm TL, west of Green Head, Western Australia, 30°00′ S, 114°27′ E, 490 m; CSIRO H 2590–11, adult male 627 mm TL, west of Leander Point, Western Australia, 29°15′ S, 113° 56′ E, 325 m; CSIRO H 822–16, immature male 541 mm TL, southwest of Shark Bay, Western Australia, 27°03′ S, 112°40′ E, 402 m; CSIRO H 2575–18, female 499 mm TL, west of Freyc�net Estuary, Western Austral�a, 26° 40′ S, 112°32′ E, 478 m; CSIRO H 6410–03, immature male 459 mm TL, west of Shark Bay, Western Austral�a, 25°31′ S, 112°10′ E, 326 m; CSIRO H 2567–10, female 448 mm TL, west of Dorre Island, Western Austral�a, 25°07′ S, 112°09′ E, 312 m; CSIRO H 1207–06, female 305 mm TL, northwest of Port Hedland, Western Australia, 18°20′ S, 117°50′ E, 430 m.Non-types. 16 spec�mens. CSIRO H 2619–10, female 650 mm TL, west of Bunbury, Western Australia, 33°22′ S, 114°31′ E, 204 m; CSIRO H 3969–15, female 600 mm TL, southwest of Fremantle, Western Australia, 33°00′ S, 114°30′ E, 324 m; CSIRO H 2591–17, adult male 610 mm TL, west of Leander Po�nt, Western Austral�a, 29°18′ S, 113°56′ E, 505 m; CSIRO H 2264–03, adult male 610 mm TL, west of Geraldton, Western Austral�a, 28°30′ S, 113°27′ E, 212 m; CSIRO H 2014–01, adult male 597 mm TL, north of Abrolhos Islands, Western Australia, 28°11′ S, 113°15′ E, 450 m; AMS I 31165–003 (2 spec�mens), female 461 mm TL and �mmature male 426 mm TL, northwest of Shark Bay, Western Austral�a, 24°00′ S, 112°09′ E, 312 m; CSIRO CA 4071, immature male 278 mm TL, southwest of Rowley Shoals, Western Australia, 18°45′ S, 117°09′ E, 356 m; CSIRO H 5786–01, female 715 mm TL, CSIRO H 5876–04, female 639 mm TL, CSIRO H 5875–05, adult male 573 mm TL,

CSIRO H 5875–06, adult male 589 mm TL, Tanjung Luar fish landing site, Lombok, Indonesia, 08°45′ S, 116°35′ E; CSIRO H 5857–09, adult male 537 mm TL, CSIRO H 5857–10, female 697 mm TL, CSIRO H 5857–11, adult male 563 mm TL, CSIRO H 5857–12, female 650 mm TL, Kedonganan fish landing site, Bali, Indonesia, 08° 45′ S, 115°10′ E.

DIAGNOSIS.— A moderate-s�zed spec�es of Squalus of the ‘m�tsukur�� group’ w�th the follow�ng comb�nat�on of characters: body moderately elongate, depth 9.9–11.5% TL; snout narrowly tr�angular, moderately long, preoral length 2.20–2.44 t�mes hor�zontal prenar�al length, 10.3–10.8% TL, mouth w�dth 1.56–1.81 (1.73) t�mes horizontal prenarial length; pre-first dorsal length 27.7–30.0 (28.5)% TL; pre-second dorsal length 60.2–62.9 (61.3)% TL; �nterdorsal space 23.5–25.6 (24.7)% TL; anterior nasal flap weakly bifurcate; first dorsal fin large, upright, first dorsal-fin height 7.0–8.0% TL; first dorsal-fin spine strong, upright and broad-based; second dorsal-fin spine long, moderately broad-based; prepectoral length 21.9–23.1 (22.6)% TL; pelv�c–caudal space 25.0–26.7 (25.9)% TL; pectoral fin of adult not falcate; pectoral-fin inner margin relatively short, 6.4–7.1% TL; caudal bar obl�que, extend�ng along base of lower lobe to caudal fork, less than 0.1–0.2 of poster�or marg�n of upper lobe, upper caudal fr�nge narrow, w�th a large dark saddle situated distinctly closer to tip of lobe than its base; flank dent�cles strongly tr�cusp�date; 43–44 monospondylous centra, 86–91 precaudal centra, 113–120 total centra; adult max�mum s�ze at least 73 cm TL.

DESCRIPTION.— Body fus�form, moderately long, nape moderately humped (most pronounced �n largest female, CSIRO H 2608–16); deepest near first dorsal-fin sp�ne (more poster�or �n smallest paratypes), max�mum depth 1.02 (0.92–1.01 �n 5 paratypes) t�mes w�dth; trunk depth 1.01 (0.87–1.01) t�mes abdomen depth; head moderately elongate 22.4 (22.5–23.1)% TL; caudal peduncle slender, 26.2 (25.0–26.7)% TL. Head not broad, w�dth 1.20 (1.14–1.24) t�mes trunk w�dth, 1.37 (1.12–1.40) t�mes abdomen w�dth; depressed forward of sp�racles, becom�ng subtr�angular �n cross-sect�on towards pectoral-fin origin; length 2.14 (2.13–2.26) in pre-vent length; he�ght 0.77 (0.71–0.76) t�mes w�dth. Snout elongate, narrowly tr�angular �n lateral v�ew, apex bluntly po�nted; lateral prenar�al marg�n not angular (sl�ghtly angular); very po�nted �n dorsal v�ew (bluntly po�nted �n some paratypes); hor�zontal length 1.52 (1.23–1.45) t�mes eye length, 0.83 (0.82–0.85) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.20 (2.25–2.44) t�mes �n preoral length. Eye narrowly oval, s�ze moderate, length 4.88 (4.03–4.63) �n head, 2.81 (2.17–2.57) t�mes he�ght; strongly notched poster�orly, notch extend�ng as a well-developed furrow half the d�stance to sp�racle (rarely reach�ng sp�racle). Sp�racle s�ze moderate, broadly crescent�c (var�able); broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.85 (3.28–4.42) �n eye length. G�ll open�ngs d�rected sl�ghtly


Figure 1. Lateral v�ew of Squalus edmundsi sp. nov. holotype (CSIRO H 2566–01, adult male 614 mm TL).

Figure 2. Ventral v�ew of the head of Squalus edmundsi sp. nov. holotype (CSIRO H 2566–01, adult male 614 mm TL).

anterodorsally from bottom to top (somet�mes upr�ght); first four subequal in size (first on left side distinctly smaller in CSIRO H 2605–05), fifth longest, height of fifth slit 2.1 (2.1–2.3)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.41 (1.33–1.38) �n preoral length; upper lab�al furrows less than tw�ce length of lower furrows; prom�nent postoral groove, longer than upper lab�al furrows, extend�ng posterolaterally from angle of jaws; �n paratypes, two ser�es of funct�onal teeth �n upper jaw, two or three ser�es �n lower. Teeth �n paratypes s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Nostr�ls small, almost transverse; anter�or nasal flap weakly bifurcate, upper lobe greatly enlarged; posterior lobe rudimentary to short (variable), flattened; �nternar�al space 2.19 (2.06–2.28) �n preoral length, 2.63 (2.58–3.06) t�mes nostr�l length. Dermal dent�cles (based on holotype) on flank very small, strongly imbricate; crowns broad, strongly tr�cusp�date w�th pronounced med�an r�dge; med�an r�dge commenc�ng well forward of

rest of crown, w�th a mes�al furrow develop�ng anter�orly and converg�ng towards poster�or t�p of crown; lateral cusps well developed, r�dges not v�s�ble (�n female paratype CSIRO H 2608–16, lateral cusps less well developed, with evidence of ridges). First dorsal fin moderate, elevated, upr�ght, broadly rounded ap�cally; anter�or marg�n very strongly convex; upper poster�or marg�n stra�ght, not vert�cal, d�rected well anterodorsally from bottom to top, weakly (rarely moderately) concave near free rear t�p; free rear t�p relat�vely th�ck basally, short; inner margin of fin almost straight; insertion of base extremely well forward of pelvic-fin origin, well posterior to free rear tip of pectoral fin; fin-spine origin above pectoral-fin insertion (slightly posterior in CSIRO H 2608–16); sp�ne base broad, exposed anter�orly well below junction of spine and soft portion of fin; soft portion of fin connected above (or near) mid-point of total spine length; sp�ne taper�ng d�stally, anter�or marg�n almost stra�ght; exposed port�on upr�ght, subequal �n length to exposed portion of second dorsal-fin spine; unexposed base only sl�ghtly longer than unexposed base of second

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dorsal-fin spine; pre-first dorsal-fin length 3.53 (3.34–3.61) times in TL; first dorsal-fin length 1.88 (1.82–2.00) t�mes �ts he�ght, 1.17 (1.11–1.25) t�mes second dorsal-fin length; first dorsal-fin height 1.67 (1.63–1.78) times second dorsal-fin height; exposed first dorsal spine length 0.65 (0.66–0.82) times height of fin. Second dorsal fin of moderate s�ze, strongly raked; anter�or marg�n convex, apex narrowly rounded; poster�or marg�n very deeply concave, max�mum concav�ty almost near m�d-po�nt of marg�n, upper port�on d�rected dorsoposter�orly strongly from bottom to top; free rear t�p elongate, �nner marg�n length 0.96 (0.93–1.04) times fin height; second dorsal-fin length 2.68 (2.60–2.80) times its height; spine length 1.11 (1.09–1.27) in height of fin; fin-spine origin well behind free rear tip of pelvic fin, exposed well below (var�able) level of junct�on w�th sp�ne and soft port�on of fin; second dorsal spine moderately broad based, slender, sharply po�nted d�stally, taper�ng rap�dly just above po�nt of exposure, sp�ne t�p extend�ng to about level of �nsert�on of fin (slightly posterior in some); interdorsal space 0.87 (0.86–0.98) �n prepectoral length, 1.11 (1.09–1.21) �n pre-first dorsal length; interdorsal groove weak. Pectoral fin small, anter�or marg�n moderately convex; �nner marg�n weakly convex, length 6.4 (6.4–7.1)% TL; apex narrowly rounded (somet�mes angular), lobe-l�ke but not falcate; poster�or marg�n moderately concave, free rear t�p broadly rounded; base very short, 2.60 (2.57–3.02) �n length of anterior margin. Pelvic fins small, anterior and posterior marg�ns almost stra�ght, apex broadly rounded, free rear t�p acute. Caudal peduncle long, taper�ng sl�ghtly to caudal fin; subcircular in cross-section anteriorly, moderately sem�c�rcular poster�orly; ventral groove well developed; lateral keels well developed, or�g�nat�ng under �nsert�on of second dorsal fin, terminating less than an eye diameter behind caudal-fin insertion; pelvic–caudal space 0.79 (0.78–0.95) �n pectoral–pelv�c space, 0.84 (0.85–0.90) �n prepectoral length; dorsal–caudal space 2.51 (2.15–2.60) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal pit moderate. Caudal fin relatively long, upper postventral marg�n moderately convex; apex of

Figure 3. Cusps of the flank denticles of Squalus edmundsi sp. nov. paratype (CSIRO H 822–16, �mmature male 541 mm TL). F�eld of v�ew w�dth 1.3 mm.

lower lobe somewhat angular; dorsal caudal marg�n 1.07 (1.07–1.14) �n head length; length of lower caudal lobe 2.00 (1.84–1.94) �n upper lobe length. Vertebral centra 113 (115–120 �n 10 paratypes), monospondylous 43 (43–44), precaudal 87 (86–91) and caudal 26 (24–30). Teeth �n upper jaw (�n paratype CSIRO H 2605–05) 12+13=25, lower jaw 11+11=22.

COLOUR.— When fresh (based on holotype): head grey�sh to grey�sh brown dorsally, paler off-wh�te to l�ght grey ventrally; l�ght and dark tonal areas sharply demarcated, �nterface extend�ng from snout to below eye, through middle of gill slits to pectoral-fin base; entire trunk mostly dark, s�m�lar to dorsal colorat�on of head. First dorsal fin dusky, free rear tip slightly paler, apical margin black-tipped; second dorsal fin dusky, apical marg�n black-t�pped, m�d- and lower poster�or marg�n pale (abruptly demarcted from black apex); dorsal sp�nes dusky, darkest anteriorly. Caudal fin mostly dusky; black upper caudal fr�nge very narrow; wh�te marg�n on upper and lower lobes of caudal fin well-demarcated; evidence of an obl�que, black�sh caudal bar along base of anter�or lower lobe, confined to notch on posterior fin margin. Pectoral and pelvic-fin upper surfaces greyish. In juvenile paratype (CSIRO H 2567–10), body colorat�on s�m�lar, w�th sl�ghtly paler ventral surface; dark marg�ns on dorsal fins more pronounced; free rear tip of first dorsal fin white (much paler than holotype), strongly contrasted

Figure 4. Lateral view of the dorsal fins of Squalus edmundsi sp. nov. paratype (CSIRO H 2605–05, female 649 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B


w�th black apex; pale area on poster�or marg�n of second dorsal fin extending almost to apex of fin, very strongly contrasted; caudal fin with a distinctive white posterior marg�n; lower caudal lobe wh�te d�stally, bordered basally with a well-defined, oblique, diffuse-edged, blackish caudal bar (not quite touching posterior margin of fin at fork); blackish upper caudal fringe better defined than in holotype; no obv�ous black marg�n on anter�or marg�n of pectoral fin; evidence of large, upper caudal blotch on central upper lobe (more pronounced �n smaller juven�le non-type, CSIRO CA 4071). In preservat�ve (based on holotype): colorat�on s�m�lar, more un�formly grey�sh, l�ght and dark tonal areas on head rema�n�ng ev�dent; pale margins on caudal fin distinct and caudal bar still ev�dent.

SIZE.— Females and males reach at least 866 (not reta�ned) and 700 mm TL (not reta�ned), respect�vely; smallest adult male 537 mm TL (non-type), �mmature males rang�ng from 278 to 541 mm TL; smallest post-natal female and male 305 and 278 mm TL, respect�vely.

DISTRIBUTION.— Upper cont�nental slope off Western Austral�a, southwest of Fremantle, Western Austral�a (ca. 33° S, 114° E) to east of Rowley Shoals, Western Austral�a (ca. 18° S, 118° E). Occurs �n depths of 204–850 m (mostly 300–500 m). Also recorded off eastern Indones�a between Kedonganan, Bal� (ca. 09° S, 115° E) and Tanjung Luar, Lombok (ca. 09° S, 116° E).

ETYMOLOGY.— Named after Matt Edmunds for h�s h�gh-qual�ty, prel�m�nary research on Austral�an Squalus undertaken as part of a summer vacat�on scholarsh�p at the CSIRO Mar�ne Laborator�es �n the early 1990’s.

VERNACULAR.— Edmunds’ Spurdog.

REMARKS.— Squalus edmundsi can be d�st�ngu�shed from all other reg�onal members of the ‘m�tsukur�� group’ �n hav�ng an obl�que caudal bar through the base of the lower caudal-fin lobe rather than a vertical bar along the posterior fin margin. It also differs in a combination of snout length, dorsal fin shape, dorsal-fin spine robustness, and other mer�st�c and morphometr�c deta�ls. It d�ffers from the morpholog�cally s�m�lar S. chloroculus, S. mitsukurii and S. montalbani, in having more upright dorsal fins with more robust spines (first dorsal-fin spine base width 1.0–1.2 vs. 0.5–0.8% TL), and a smaller max�mum s�ze (males atta�n�ng 70 cm TL vs. >85 cm TL �n the other three spec�es). Squalus edmundsi d�ffers from the three superficially-similar species belonging to the ‘highfin megalops group’, S. albifrons, and S. altipinnis (Last, Wh�te and Stevens, 2007; Part 5 of th�s �ssue) �n hav�ng a dark caudal bar that e�ther extends or almost reaches to the caudal-fin posterior margin at the caudal fork vs. an ent�rely pale poster�or marg�n, and these spec�es and S. notocaudatus (Last, Wh�te and Stevens, 2007; Part 5 of th�s �ssue) by a longer, more po�nted snout (preoral snout 10.3–10.8 vs. 8.4–9.5% TL; prenar�al length 5.6–5.8 vs. 4.3–5.0% TL).

Squalus grahami sp. nov.


Squalus sp. F: Last and Stevens, 1994, Sharks and Rays of Australia, pp 49, 97, fig. 8.26, 8.36, pl. 5; Compagno, Dando and Fowler, 2005, A Field Guide to the Sharks of the World, p 81, fig, pl 2.

Holotype. CSIRO H 4476–01, adult male 602 mm TL, northeast of Batemans Bay, New South Wales, 35°29′ S, 150°46′ E, 234 m, 4 December 1996.Paratypes. 30 spec�mens. CSIRO H 1347–01, female 380 mm TL, east of Fl�nders Reefs, Queensland, 17° 29′ S, 149°32′ E, 504 m; CSIRO H 1406–01, female 221 mm TL, Queensland Trough, east of H�nch�nbrook Island, Queensland, 17°55′ S, 147°06′ E, 402 m; CSIRO H 453, adult male 526 mm TL, CSIRO H 457–01, female 623 mm TL, northeast of H�nch�nbrook Island, Queensland, 17°57′ S, 147°03′ E, 300 m; CSIRO H 2468–02, female 653 mm TL, east of H�nch�nbrook Island, Queensland, 17°57′ S, 147°02′ E, 306 m; CSIRO H 2469–04, female 649 mm TL, CSIRO H 2469–05, female 631 mm TL, east of H�nch�nbrook Island, Queensland, 17°58′ S, 147°01′ E, 264 m; CSIRO H 454, adult male 550 mm TL, CSIRO H 455, female 635 mm TL, CSIRO H 456, adult male 550 mm TL, Townsv�lle Trough, Queensland, 17°58′ S, 147°00′ E, 220 m; CSIRO H 1311–03, female 653 mm TL, Mar�an Plateau, east of Whitsunday Islands, Queensland, 19°29′ S, 150°17′ E, 328 m; CSIRO H 1346–01, �mmature male 294 mm TL, North East Slope, Queensland; CSIRO H 602–02, female 367 mm TL, CSIRO H 1312–06, female 286 mm TL, CSIRO H 1312–07, female 340 mm TL, CSIRO H 1312–08, female 282 mm TL, CSIRO H 1312–09, �mmature

Figure 5. Juvenile coloration of the caudal fin of Squalus edmundsi sp. nov. (CSIRO CA 4071, �mmature male 278 mm TL).

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male 279 mm TL, south of Saumarez Reef, Queensland, 22°42′ S, 154°05′ E, 419 m; CSIRO H 644–03, adult male 520 mm TL, CSIRO H 644–05, adult male 531 mm TL, CSIRO H 644–06, adult male 523 mm TL, CSIRO H 1344–01, female 232 mm TL, CSIRO H 1344–02, juven�le male 238 mm TL, CSIRO H 1344–03, �mmature male 327 mm TL, south of Saumarez Reef, Queensland, 22°49′ S, 154°10′ E, 450 m; CSIRO H 452, female 624 mm TL, southwest of Saumarez Reef, Queensland, 22°52′ S, 152°42′ E, 225 m; CSIRO H 2688–02, adult male 572 mm TL, east of Coffs Harbour, New South Wales, 30°24′ S, 153°23′ E, 148 m; CSIRO H 4682–01, adult male 558 mm TL, CSIRO H 4682–02, female 689 mm TL, CSIRO H 4682–03, female 702 mm TL, CSIRO H 4682–04, adult male 571 mm TL, east of Broken Bay, New South Wales, 33°32′ S, 152°00′ E, 329 m; CSIRO H 4476–08, female 578 mm TL, collected w�th holotype.Non-types.— 18 spec�mens. CSIRO H 5635–13, female 643 mm TL, east of Rock�ngham Bay, Queensland, 18°08′ S, 147°09′ E, 248 m; CSIRO H 602–13, female 355 mm TL, south of Saumarez Reef, Queensland, 22° 42′ S, 154°05′ E, 419 m; CSIRO H 4623–03, female 607 mm TL, east of Terr�gal, New South Wales, 33° 26′ S, 152°06′ E, 383 m; NMV A 29562–001, female 675 mm TL, east of Sydney, New South Wales, 33°28′ S, 152°00′ E, 230 m; CSIRO H 4476–02, female 595 mm TL, CSIRO H 4476–03, adult male 522 mm TL, CSIRO H 4476–04, adult male 525 mm TL, CSIRO H 4476–05, female 580 mm TL, CSIRO H 4476–06, female 592 mm TL, CSIRO H 4476–07, female 552 mm TL, CSIRO H 4476–09, female 594 mm TL, CSIRO H 4476–10, female 556 mm TL, CSIRO H 4476–11, adult male 542 mm TL, collected w�th holotype; CSIRO H 4771–01, adult male 516 mm TL, east of Brush Island, New South Wales, 35°30′ S, 150°45′ E, 333 m; CSIRO H 4477–01, female 663 mm TL, CSIRO H 4477–02, female 598 mm TL, CSIRO H 4709–02, female 711 mm TL, east of Brush Island, New South Wales, 35°34′ S, 150°44′ E, 388 m; CSIRO H 4708–01, �mmature male 549 mm TL, east of Batemans Bay, New South Wales, 35°48′ S, 150°34′ E, 291 m.

DIAGNOSIS.— A moderate-s�zed spec�es of Squalus of the ‘m�tsukur�� group’ w�th the follow�ng comb�nat�on of characters: body very elongate, depth 10.1–12.6% TL; snout narrow, moderately long, preoral length 2.38–2.53 t�mes hor�zontal prenar�al length, 10.5–11.3% TL, mouth w�dth 1.54–1.77 (1.66) t�mes hor�zontal prenar�al length; pre-first dorsal length 28.9–31.3 (30.0)% TL; pre-second dorsal length 60.2–62.7 (61.2)% TL; �nterdorsal space 22.3–24.7 (23.8)% TL; dorsal fins small, raked, first dorsal-fin height 6.3–7.2% TL; first dorsal-fin spine short, weak; second dorsal-fin spine slender with moderately broad base; prepectoral length 21.7–23.6 (22.4)% TL; pelv�c–caudal space 24.0–26.5 (25.4)% TL; pectoral fin of adult slightly falcate; pectoral-fin inner marg�n relat�vely short, 7.1–7.8% TL; caudal bar almost upr�ght, extend�ng narrowly from the caudal fork up the

poster�or marg�n of the upper lobe for usually about 0.4 (rarely to 0.6) of �ts length �n �mmatures, upper caudal fr�nge narrow, somet�mes w�th a narrow central blotch on upper lobe; flank denticles weakly tricuspidate; 37–42 monospondylous centra, 80–87 precaudal centra, 105–116 total centra; adult max�mum s�ze up to 71 cm TL.

DESCRIPTION.— Body fus�form, elongate (more robust �n large female paratypes), nape sl�ghtly humped (more pronounced �n largest paratypes); deepest near first dorsal-fin spine, maximum depth 0.94 (0.88–1.00 �n paratypes) t�mes w�dth; trunk depth 0.97 (0.86–0.98) t�mes abdomen depth; head short 22.0 (22.6–24.0)% TL; caudal peduncle moderately slender, 26.3 (24.0–26.5)% TL. Head somewhat narrow, w�dth 1.08 (1.05–1.14) t�mes trunk w�dth, 1.25 (1.10–1.28) t�mes abdomen w�dth; depressed forward of sp�racles, becom�ng subtr�angular in cross-section towards pectoral-fin origin; length 2.15 (2.07–2.21) �n pre-vent length; he�ght 0.82 (0.74–0.84) t�mes w�dth. Snout long, narrowly tr�angular �n lateral v�ew, apex bluntly po�nted; lateral prenar�al marg�n somewhat angular; po�nted �n dorsal v�ew; hor�zontal length 1.59 (1.46–1.56) t�mes eye length, 1.00 (0.90–1.01) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.38 (2.43–2.53) t�mes �n preoral length. Eye oval, s�ze moderately large, length 4.47 (4.21–4.68) �n head, 2.55 (2.24–2.58) t�mes he�ght; strongly notched poster�orly, notch extend�ng as a well-developed furrow to anteroventral marg�n of sp�racle (usually more pronounced anter�orly, somet�mes barely extend�ng to sp�racle). Sp�racle s�ze small to moderate, broadly (somet�mes narrowly) crescent�c; broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.89 (3.33–4.11) in eye length. Gill openings almost upright; first four subequal in size, fifth longest, height of fifth slit 2.1 (1.9–2.3)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.55 (1.43–1.53) �n preoral length; upper lab�al furrows about tw�ce length of lower furrows (or less); prom�nent postoral groove, much longer than upper lab�al furrows (usually longer �n paratypes), extend�ng posterolaterally from angle of jaws; two ser�es of funct�onal teeth �n upper jaw, two ser�es �n lower. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke; cusps d�rected strongly laterally, low; base of tooth broader than length of cusp. Nostr�ls small, almost transverse; anterior nasal flap strongly b�furcate, upper lobe largest, broadest; poster�or lobe narrow, thallate (var�able �n paratypes); �nternar�al space 2.23 (2.11–2.28) �n preoral length, 2.74 (2.67–3.04) t�mes nostr�l length. Dermal dent�cles (based on the holotype) on flank very small; crown weakly tricuspidate with pronounced med�an r�dge, med�an cusp apex broadly tr�angular; med�an r�dge or�g�n sl�ghtly anter�or to rest of crown; r�dge w�th mes�al furrow converg�ng towards poster�or t�p of crown; lateral cusps weakly developed, blunt to rud�mentary, d�stal r�dges not ev�dent. F�rst dorsal fin somewhat raked, broadly rounded apically; anterior marg�n weakly convex; upper poster�or marg�n weakly convex, not vert�cal, d�rected sl�ghtly anterodorsally from


Figure 6. Lateral v�ew of Squalus grahami sp. nov. holotype (CSIRO H 4476–01, adult male 602 mm TL).

Figure 7. Ventral v�ew of the head of Squalus grahami sp. nov. holotype (CSIRO H 4476–01, adult male 602 mm TL).

bottom to top, moderately concave near free rear t�p; free rear tip moderately thick basally, short; inner margin of fin almost stra�ght; �nsert�on of base extremely well forward of pelvic-fin origin, well posterior to free rear tip of pectoral fin; fin-spine posterior to pectoral-fin insertion; sp�ne base narrow, exposed anter�orly below junct�on of spine and soft portion of fin; soft portion of fin connected sl�ghtly above m�d-po�nt of total sp�ne length (�n paratype CSIRO H 4682–03 close to sp�ne apex); sp�ne taper�ng d�stally, anter�or marg�n almost stra�ght; exposed port�on of sp�ne raked, shorter than exposed port�on of second dorsal-fin spine; pre-first dorsal length 3.44 (3.19–3.46) times in TL; first dorsal-fin length 1.97 (2.04–2.28) t�mes �ts he�ght, 1.12 (1.09–1.19) t�mes second dorsal-fin length; first dorsal-fin height 1.50 (1.25–1.66) times second dorsal-fin height; exposed first dorsal spine length 0.56 (0.49–0.67) times height of fin. Second dorsal fin

of moderate s�ze, strongly raked; anter�or marg�n sl�ghtly convex, apex narrowly rounded; poster�or marg�n deeply concave, max�mum concav�ty well below m�d-po�nt of marg�n, upper port�on d�rected sl�ghtly dorsoposter�orly from bottom to top; free rear t�p very elongate, �nner margin length 1.15 (0.98–1.18) times fin height; second dorsal-fin length 2.64 (2.53–3.08) times its height; spine length 1.03 (0.86–1.14) in height of fin; fin-spine origin well behind free rear tip of pelvic fin, usually exposed at level of junction with spine and soft portion of fin; second dorsal sp�ne moderately broad based (more pronounced �n largest paratypes), slender, anter�or marg�n sl�ghtly convex, weakly recurved d�stally, apex sharply po�nted, taper�ng just above po�nt of exposure; sp�ne t�p extend�ng behind level of insertion of fin (over in some paratypes); �nterdorsal space 0.88 (0.91–1.06) �n pre-pectoral length, 1.18 (1.22–1.39) in pre-first dorsal length; interdorsal

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Table 1. Proport�onal d�mens�ons as percentages of total length for the holotypes and 5 paratypes of Squalus edmundsi sp. nov. (holotype CSIRO H 2566–01) and Squalus grahami sp. nov. (holotype CSIRO H 4476–01).

S. edmundsi sp. nov. S. grahami sp. nov. Holotype Paratypes Holotype Paratypes

M�n. Max. M�n. Max.TL – Total length 614 508 738 602 558 702PCL – Precaudal length 78.8 78.4 79.8 78.2 77.8 80.5PD2 – Pre-second dorsal length 61.1 60.2 62.9 59.1 58.8 62.0PD1 – Pre-first dorsal length 28.4 27.7 30.0 28.4 28.6 30.5SVL – Pre-vent length 47.9 48.8 51.6 47.3 47.8 50.9PP2 – Prepelv�c length 45.9 47.0 49.5 45.3 45.5 50.1PP1 – Prepectoral length 22.1 21.9 23.1 21.7 22.1 23.6HDL – Head length 22.4 22.5 23.1 22.0 22.7 24.0PG1 – Prebranch�al length 19.2 19.3 19.9 19.0 19.5 20.5PSP – Presp�racular length 13.3 13.2 13.6 12.9 12.9 14.0POB – Preorb�tal length 8.0 7.9 8.1 8.3 8.3 9.0PRN – Prenar�al length 5.8 5.6 5.8 5.6 5.7 6.3POR – Preoral length 10.3 10.4 10.8 10.5 10.8 11.3INLF – Inner nostr�l-lab�al furrow space 4.8 4.5 5.0 4.7 4.5 5.0MOW – Mouth w�dth 7.3 7.8 7.9 6.8 7.1 7.9ULA – Lab�al furrow length 2.2 2.0 2.3 2.2 2.3 2.8INW – Internar�al space 4.7 4.6 5.0 4.7 4.7 5.4INO – Interorb�tal space 8.4 8.2 8.8 7.8 7.6 8.5EYL – Eye length 4.6 4.9 5.7 4.9 4.9 5.5EYH – Eye he�ght 1.6 2.0 2.3 1.9 1.9 2.3SPL – Sp�racle length 1.2 1.2 1.5 1.5 1.3 1.7GS1 – F�rst g�ll-sl�t he�ght 1.6 1.4 2.0 1.5 1.4 1.8GS5 – F�fth g�ll-sl�t he�ght 2.1 2.1 2.3 2.1 1.9 2.3IDS – Interdorsal space 25.5 23.5 25.6 23.7 20.6 23.8DCS – Dorsal-caudal space 10.2 9.6 10.9 10.8 9.9 11.1PPS – Pectoral-pelv�c space 20.7 20.8 23.8 20.6 20.3 24.8PCA – Pelv�c-caudal space 26.2 25.0 26.7 26.3 24.0 26.5D1L – F�rst dorsal length 14.6 13.8 14.8 14.1 14.8 15.2D1A – F�rst dorsal anter�or marg�n 11.9 11.3 12.8 11.7 12.0 13.0D1B – F�rst dorsal base length 9.3 8.3 9.6 8.6 9.1 10.1D1H – F�rst dorsal he�ght 7.7 7.0 8.0 7.0 6.3 7.2D1I – F�rst dorsal �nner marg�n 5.5 4.9 5.6 5.9 5.1 6.0D1P – F�rst dorsal poster�or marg�n 9.2 8.5 9.1 8.0 7.5 8.6D1ES – F�rst dorsal sp�ne length 5.0 4.8 5.7 3.9 3.1 4.2D1BS – F�rst dorsal sp�ne base w�dth 1.2 1.0 1.2 0.6 0.6 0.8D2L – Second dorsal length 12.5 11.6 13.0 13.9 12.4 14.4D2A – Second dorsal anter�or marg�n 11.0 10.2 11.8 11.2 10.2 12.2D2B – Second dorsal base length 8.1 7.5 8.6 8.5 8.0 9.6D2H – Second dorsal he�ght 4.6 4.2 4.9 4.6 4.0 5.1D2I – Second dorsal �nner marg�n 4.5 4.1 4.6 5.3 4.3 5.1D2P – Second dorsal poster�or marg�n 5.0 4.4 5.0 5.6 4.8 6.2D2ES – Second dorsal sp�ne length 5.2 5.0 5.5 4.8 3.5 5.4D2BS – Second dorsal sp�ne base w�dth 0.8 0.8 1.0 0.7 0.7 0.8


Table 1. cont’d.

S. edmundsi sp. nov. S. grahami sp. nov. Holotype Paratypes Holotype Paratypes

M�n. Max. M�n. Max.P1A – Pectoral anter�or marg�n 14.0 14.3 15.1 14.4 14.1 15.0P1I – Pectoral �nner marg�n 6.4 6.4 7.1 7.5 7.1 7.8P1B – Pectoral base length 5.4 4.9 5.9 5.0 5.3 6.1P1P – Pectoral poster�or marg�n 9.9 10.0 11.2 10.0 10.6 12.0P2L – Pelv�c length 10.4 8.9 10.4 11.2 10.5 11.5P2H – Pelv�c he�ght 4.7 3.9 5.1 3.9 4.1 4.6P2I – Pelv�c �nner marg�n 5.5 3.7 5.6 5.7 5.0 6.4CDM – Dorsal caudal marg�n 21.0 20.0 21.1 21.7 19.2 22.0CPV – Preventral caudal marg�n 10.5 10.8 11.4 11.9 11.0 12.1CPU – Upper postventral caudal marg�n 15.7 15.0 15.9 16.4 14.2 16.5CPL – Lower postventral caudal marg�n 4.3 3.6 5.0 5.1 4.0 5.0CFW – Caudal fork w�dth 6.5 6.2 6.9 6.3 6.2 7.2CFL – Caudal fork length 8.6 8.3 9.2 9.2 9.3 10.0HANW – Head w�dth at nostr�ls 6.8 6.8 7.3 8.2 8.1 8.9HAMW – Head w�dth at mouth 10.8 11.1 11.7 10.3 11.2 11.5HDW – Head w�dth 12.0 11.8 13.3 11.2 11.4 13.2TRW – Trunk w�dth 10.0 9.8 11.7 10.4 10.3 12.1ABW – Abdomen w�dth 8.7 8.4 11.9 9.0 9.5 11.7TAW – Ta�l w�dth 6.4 5.7 6.7 5.8 6.5 7.1CPW – Caudal peduncle w�dth 3.0 2.4 3.0 2.8 2.9 3.2HDH – Head he�ght 9.3 8.3 10.0 9.2 9.1 10.3TRH – Trunk he�ght 10.2 9.2 10.7 9.8 9.8 11.5ABH – Abdomen he�ght 10.1 9.9 11.5 10.1 10.5 12.6TAH – Ta�l he�ght 6.6 6.2 6.6 6.0 6.2 7.3CPH – Caudal peduncle he�ght 2.3 2.2 2.4 2.4 2.3 2.6CLO – Clasper outer length 3.4 3.0 3.0 4.0 4.0 4.0CLI – Clasper �nner length 6.2 6.1 6.1 7.2 7.7 8.2CLB – Clasper base w�dth 1.1 1.2 1.2 1.6 1.3 1.6

ridge obvious (variable in paratypes). Pectoral fin small, length 7.5 (7.1–7.8)% TL; anter�or marg�n weakly convex, apex narrowly rounded, sl�ghtly falcate; poster�or marg�n moderately concave, �nner marg�n convex, free rear tip bluntly angular; fin base very short, 2.89 (2.46–2.83) in length of anterior margin. Pelvic fins moderate, anter�or and poster�or marg�ns almost stra�ght, apex broadly rounded, free rear t�p acute. Caudal peduncle long, tapering very slightly to caudal fin; subcircular in cross-sect�on anter�orly, broadly sem�c�rcular poster�orly; ventral groove well developed (var�able �n paratypes); lateral keels well developed, or�g�nat�ng under �nsert�on of second dorsal fin (or slightly behind), terminating less than an eye diameter behind caudal-fin insertion; pelvic–caudal space 0.78 (0.76–1.03) �n pectoral–pelv�c space, 0.83 (0.85–0.94) �n prepectoral length; dorsal–caudal space 2.28 (2.04–2.43) �n �nterdorsal length; dorsal

caudal p�t well developed, ventral caudal p�t rud�mentary. Caudal fin moderate, posterior margin of upper lobe very strongly convex, base broad; apex of lower lobe narrowly rounded to somewhat angular; dorsal caudal marg�n 1.01 (1.03–1.21) �n head length; length of lower caudal lobe 1.83 (1.64–1.81) �n upper lobe length. Vertebral centra 113 (108–116), monospondylous 40 (38–42), precaudal 86 (80–87) and caudal 27 (26–32). Teeth �n upper jaw (�n paratype CSIRO H 2469–05) 13+14=27, lower jaw 12+11=23.

COLOUR.— When fresh (based on holotype): dorsal surface dark grey, off-wh�te to grey�sh below; l�ght and dark tonal areas well demarcated on head, extend�ng from snout to below eye, through m�ddle of g�ll sl�ts to pectoral-fin base; not obviously demarcated on belly; tail almost un�formly grey�sh above and below. F�rst dorsal

80

fin almost uniformly dark grey, base slightly paler, narrow black apical margin; second dorsal fin uniformly greyish, no evidence of darker markings; dorsal-fin spines dusky. Caudal bar black�sh, pronounced, extend�ng dorsally from fork, cover�ng basal half of upper lobe poster�orly; d�stal half of poster�or marg�n of upper lobe pale, rema�nder of lobe dusky; ventral lobe almost un�formly pale; fleshy portion of fin bordered above and below by dark, d�agonal, grey�sh caudal str�pes. Small paratype (CSIRO H 602–02) significantly paler than holotype, greyish above, almost wh�te below; l�ght and dark tonal areas less strongly demarcated w�th more pronounced dark marg�ns along dorsal fins; a broad dark edge on first dorsal fin extending from upper anterior margin along fin towards free rear tip; second dorsal fin similar, dark marking very broad, not extend�ng onto free rear t�p; dark caudal bar very d�st�nct, strongly demarcated from wh�te areas on upper and lower lobes; upper caudal fr�nge form�ng a well-developed saddle on m�ddle of lobe, fa�nt upper caudal blotch on m�d-outer lobe; upper caudal str�pe narrow, lower str�pe appear�ng as a prom�nent black basal blotch; narrow black anterior margin on pectoral fin. In preservat�ve (based on holotype): narrow dark marg�ns on dorsal fins more evident than in fresh state; dorsal surface of pectoral fin similar to upper body, posterior marg�ns only sl�ghtly paler; ventral surface of pectoral fins dusky with a whitish base; upper pelvic fins greyish w�th broad, pale poster�or marg�n, more un�formly pale ventrally. Neonatal paratype (CSIRO H 1406–01) s�m�lar to fresh paratype (CSIRO H 602–02), more yellow�sh, dark coloration on fins similarly disposed; upper caudal blotch and upper caudal str�pe prom�nent.

SIZE.— Females and males reach at least 711 (non-type) and 602 mm TL (holotype), respect�vely; smallest adult male 516 mm TL, �mmature males rang�ng from 238 to 327 mm TL; smallest post-natal spec�men 232 mm TL. Graham (2005) prov�des bas�c b�olog�cal data for several Squalus spec�es �nclud�ng S. grahami (as S. sp. F) and reports max�mum s�zes of 730 and 620 mm TL for

Figure 8. Cusps of the flank denticles of Squalus grahami sp. nov. (CSIRO H 602–02, female 367 mm TL). F�eld of v�ew w�dth 0.9 mm.

females and males, respect�vely.

DISTRIBUTION.— Upper cont�nental slope off eastern Austral�a, from north of Brush Island, New South Wales (ca. 35°S, 150°E) to Cape York, Queensland (ca. 10°S, 144°E). Known from depths of 148–504 m but trawled ma�nly �n depths of 220–450 m.

ETYMOLOGY.— Named after prom�nent Austral�an

A

B

Figure 9. Lateral view of the dorsal fins of Squalus grahami sp. nov. holotype (CSIRO H 4476–01, adult male 602 mm TL) – A. first dorsal fin, B. second dorsal fin.

Figure 10. Juvenile coloration of the caudal fin of Squalus grahami sp. nov. (CSIRO H 1406–01 female 221 mm TL).


�chthyolog�st, Ken Graham (NSW Department of Pr�mary Industr�es, Cronulla), who has contr�buted greatly to our knowledge of the sharks and rays of southeastern Austral�a, and who collected the pr�mary type, as well as the major�ty of New South Wales mater�al of th�s spec�es.

VERNACULAR.— Eastern Longnose Spurdog.

REMARKS.— Squalus grahami can be d�st�ngu�shed from all other members of the ‘m�tsukur�� group’ by the comb�nat�on of a narrow, moderately long snout, small raked dorsal fins, small dorsal-fin spines, and other mer�st�c and morphometr�c deta�ls. It d�ffers from the morpholog�cally s�m�lar S. montalbani, S. chloroculus and S. mitsukurii �n hav�ng a much smaller max�mum s�ze (ca. 600 vs. >850 mm TL for males), a sl�ghtly longer snout (preoral length 10.5–11.3 vs. 9.5–10.1, 9.3–10.6 and 9.4–10.8% TL respect�vely), and fewer monospondylous centra (37–42 vs. 41–47, 43–46 and 45–51 respect�vely). Squalus grahami d�ffers from S. edmundsi �n hav�ng an upr�ght caudal bar (rather than obl�que), more strongly raked dorsal fins with short, narrow spines (exposed first dorsal sp�ne length 3.1–4.2 vs. 4.8–5.7 TL), a sl�ghtly longer snout (preorb�tal length 8.3–9.0 vs. 7.9–8.1% TL), and sl�ghtly less precaudal centra (80–87 vs. 86–92).

ACKNOWLEDGEMENTS

The new species were first identified in a study of the Austral�an chondr�chthyan fauna (Last and Stevens, 1994) that was jo�ntly funded by the CSIRO and the F�sher�es Research and Development Corporat�on (FRDC). S�ncere thanks go to Alasta�r Graham for assembl�ng data on Austral�an mater�al, Lou�se Conboy for prepar�ng �mages of the types, T�m Founta�n for prepar�ng rad�ographs and obta�n�ng mer�st�c �nformat�on, Tony Rees for h�s ass�stance w�th prepar�ng SEMs, Matt Edmunds for h�s prel�m�nary research on th�s group, Ken Graham for supply�ng fresh collect�on mater�al, and Gordon Yearsley and John Pogonosk� for other techn�cal support and ed�tor�al ass�stance. Spec�al thanks also go to curators and curator�al staff that ass�sted �n prov�d�ng research mater�al and reg�strat�on �nformat�on: Mark McGrouther (AMS) and Mart�n Gomon and D� Bray

(NMV). Indones�an mater�al was collected as part of a 5-year ACIAR funded project and �mportant contr�butors to th�s project �nclude Junaed� (Kedonganan F�sh Market, Bali), processors and fishers at Tanjung Luar fish market (Lombok), Fahm� (LIPI), Dharmad� (RCCF), Jenny G�les (QDPI&F), Ono Kurnaen Sumadh�harga (LIPI), Subhat Nurhak�m (RCCF), Ian Potter (Murdoch Un�vers�ty), and Cathy D�chmont and Steve Blaber (CSIRO).

REFERENCES


Graham, K.J. (2005) D�str�but�on, populat�on structure and b�olog�cal aspects of Squalus spp. (Chondr�chthyes: Squal�formes) from New South Wales and adjacent Austral�an waters. Marine and Freshwater Research, 56, 405–416.


Last, P.R., Wh�te, W.T. and Stevens, J.D. (2007) Part 5 — New spec�es of Squalus of the ‘highfin megalops group’ from the Australas�an reg�on, p. 39–53. In: Descr�pt�ons of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last P.R., Wh�te W.T. and Pogonosk� J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.




Part 8 — Squalus nasutus sp. nov., a new long-snout spurdog of the ‘japonicus group’ from the Indian Ocean

Peter R. Last, L�ndsay J. Marshall and W�ll�am T. Wh�te


ABSTRACT.— A new spec�es of spurdog, Squalus nasutus sp. nov., �s descr�bed based on spec�mens from the upper cont�nental slope off Western Austral�a. The new spec�es, wh�ch has been confused w�th S. japonicus from the western North Pacific, belongs to a subgroup of Squalus whose members all have a very elongate snout. Squalus nasutus differs from its closely related congeners in a combination of fin coloration, meristics and morphometrics of the head, trunk and pectoral fins. It appears to occur outside Australian seas and intraspecific variation between populations across the Indo–West Pacific is discussed.

Key words. Squalo�dea – Squal�dae – Squalus nasutus – new species – Indo–West Pacific


INTRODUCTION

Last and Stevens (1994) identified 9 Squalus spec�es from Austral�an waters of wh�ch 6 appear to be formally undescr�bed. One of these spec�es belongs to the ‘japon�cus group’. The group presently conta�ns three nom�nal spec�es, S. japonicus Ish�kawa, 1908 from the western North Pacific, and S. melanurus Fourmano�r and R�vaton, 1979 and S. rancureli Fourmano�r and R�vaton, 1979 from New Caledon�a. These spec�es are character�sed by a very slender body and a very long snout. A new spec�es of the ‘japonicus group’, informally identified as Squalus sp. E (sensu Last and Stevens, 1994), �s descr�bed based on Austral�an mater�al and compar�sons are made w�th s�m�lar forms from the Ind�an Ocean.

METHODS

Methods follow those outl�ned �n Part 1 of th�s �ssue (Last et al., 2007). Morphometr�cs and mer�st�cs were taken from the holotype (CSIRO H 2590–12) and the follow�ng 5 paratypes: CSIRO H 1652–02, CSIRO H 2032–01, CSIRO H 2567–08, CSIRO H 2598–07 and CSIRO H 2898–07 (Table 1). In add�t�on, mer�st�cs were taken from the follow�ng 13 paratypes: CSIRO CA 3290, CSIRO CA 4055, CSIRO CA 4110, CSIRO H 1207–07, CSIRO H 1207–08, CSIRO 1652–01, CSIRO H 1693–01, CSIRO H 1693–02, CSIRO H 1694–01, CSIRO H 2032–02, CSIRO H 2608–15 and WAM P 28086–006 (2 spec�mens). Shape data was obta�ned from an add�t�onal 4 spec�mens from Indones�a and 3 spec�mens from the Ph�l�pp�nes of a’ japon�cus group’ Squalus thought to be conspecific with the new species. A subset of morphometric characters poss�bly useful �n d�scr�m�nat�ng members of

the ‘japon�cus group’ were obta�ned from the rema�n�ng post-juven�le paratypes. In the formal descr�pt�on, morphometr�c and mer�st�c values for the holotype are given first followed in parentheses by the ranges of the paratypes. Mer�st�cs were taken from rad�ographs of 17 Austral�an, 4 Indones�an and 5 Ph�l�pp�ne spec�mens of the new spec�es as well as 4 spec�mens of S. japonicus. Type spec�mens are depos�ted �n the Austral�an Nat�onal F�sh Collect�on, Hobart, Austral�a (CSIRO) and Western Austral�an Museum, Perth (WAM). Other mater�al used �n th�s study are depos�ted �n the �chthyolog�cal collect�ons of the Austral�an Museum, Sydney (AMS), Nat�onal Museum, V�ctor�a (NMV), S�ll�man Un�vers�ty Mar�ne Laboratory, Ph�l�pp�nes (SUML), Un�vers�ty of Flor�da, USA (UF) and Hokka�do Un�vers�ty, Japan (HUMZ); their registration numbers are prefixed with these acronyms.

Squalus nasutus sp. nov.

F�gs 1a, 2a, 3, 4, 5a; Table 1

Squalus sp. E: Last and Stevens, 1994, Sharks and Rays of Australia, pp 49, 96, fig. 8.25, 8.36, pl. 5; Compagno, Dando and Fowler, 2005, Sharks of the World, p 80–1, figs, pl. 2; Wh�te et al., 2006, Economically Important Sharks and Rays of Indonesia, pp 74–75.

Holotype. CSIRO H 2590–12, female 508 mm TL, west of Leander Point, Western Australia, 29°15′ S, 113°56′ E, 325 m, 6 Feb 1991.Paratypes. 18 spec�mens. CSIRO H 2608–15, female 411 mm TL, Rottnest Canyon, Western Austral�a, 31° 57′ S, 115°08′ E, 850 m; CSIRO H 2598–07, female

84

465 mm TL, west of Green Head, Western Austral�a, 29°58′ S, 114°27′ E, 380 m; CSIRO H 2567–08, adult male 470 mm TL, west of Dorre Island, Western Austral�a, 25°09′ S, 112°09′ E, 312 m; CSIRO 1652–01, female 315 mm TL, CSIRO H 1652–02, female 459 mm TL, northwest of Port Hedland, Western Australia, 18°25′ S, 117°48′ E, 375 m; CSIRO H 1207–07, female 537 mm TL, CSIRO H 1207–08, female 496 mm TL, northwest of Port Hedland, Western Australia, 18°20′ S, 117°50′ E, 430 m; CSIRO CA 3290, female 549 mm TL, southwest of Rowley Shoals, Western Australia, 18°10′ S, 118°20′ E, 300 m; CSIRO CA 4055, female 527 mm TL, southwest of Rowley Shoals, Western Australia, 18°11′ S, 118° 04′ E, 400 m; CSIRO H 2898–07, adolescent male 452 mm TL, north-northwest of Port Hedland, Western Australia, 18°07′ S, 118°12′ E, 361 m; WAM P 28086–006 (2 spec�mens), female 452 mm TL and �mmature male 380 mm TL, Rowley Shoals, Western Australia, 17°49′ S, 118°41′ E, 310 m; CSIRO CA 4110, adult male 497 mm TL, east of Rowley Shoals, Western Australia, 17°18′ S, 120°09′ E, 305 m; CSIRO H 1693–01, immature male 361 mm TL, CSIRO H 1693–02, female 306 mm TL, Rowley Shoals, Western Australia, 17°02′ S, 120°05′ E, 409 m; CSIRO H 1694–01, female 425 mm TL, Rowley Shoals, Western Australia, 16°57′ S, 120°14′ E, 413 m; CSIRO H 2032–01, adult male 461 mm TL, CSIRO H 2032–02, female 404 mm TL, northeast of Merma�d Reef, Rowley Shoals, Western Australia, 16°54′ S, 120°25′ E, 392 m.Non-types. 17 spec�mens. CSIRO H 6413–01, female 590 mm TL, west of Shark Bay, Western Austral�a, 25° 03′ S, 112°08′ E, 340 m; AMS I 43986–001, female 570 mm TL, NMV A 29560–001, female 531 mm TL, WAM P 32841–001 (2 spec�mens), adult male 499 mm TL and female 472 mm TL, northwest of Shark Bay, Western Australia, 23°40′ S, 112°47′ E, 337 m; CSIRO H 6125–04, adult male 468 mm TL, Kedonganan fish landing site, Bali, Indonesia, 08°45′ S, 115°10′ E; CSIRO H 5860–01, female 546 mm TL, CSIRO H 5860–02, female 546 mm TL, CSIRO H 5860–03, female 549 mm TL, CSIRO H 6484–01, female 575 mm TL, WAM P 32842–001, female 520 mm TL, Cilacap fish landing site, Central Java, Indonesia, 07°40′ S, 109°00′ E; CSIRO H 4132–02, male 473 mm TL, CSIRO H 4132–03, female 479 mm TL, CSIRO H 4132–04, female 541 mm TL, SUML F 1131, adult male 443 mm TL, SUML F 1151, female 486 mm TL, Bol�nao even�ng market, Ph�l�pp�nes.

DIAGNOSIS.— A small spec�es of Squalus of the ‘japon�cus group’ w�th the follow�ng comb�nat�on of characters: body very elongate, depth 9.4–11.4% TL; snout narrow, long, preoral length 1.85–2.01 t�mes hor�zontal prenar�al length, 11.1–11.7% TL; eye large, 4.3–5.1% TL; secondary lobe of anterior nasal flap well developed; dorsal fins small, raked; first dorsal-fin spine short, weak; second dorsal-fin spine short with strong base; pectoral fin of adult not falcate; pectoral-fin inner marg�n short, 6.6–7.1% TL; caudal bar almost upr�ght, short, broad, extend�ng from the caudal fork up the

poster�or marg�n of the upper lobe for 0.4–0.5 of �ts length, upper caudal blotch somewhat l�near, located near dorsal margin of fin lobe; flank denticles weakly tricuspidate; 36–39 monospondylous centra, 78–81 precaudal centra, 103–109 total centra; adult s�ze at least 60 cm TL.

DESCRIPTION.— Body elongate fus�form to sl�ghtly compressed (bulg�ng marg�nally at belly, max�mum depth 1.03 (1.04–1.13) t�mes w�dth); head long 23.6 (23.0–24.1 �n paratypes)% TL; caudal peduncle elongate, narrow, depressed sl�ghtly 24.9 (25.2–26.6)% TL. Head very depressed forward of sp�racles, decreas�ngly less so towards pectoral or�g�n, length 2.19 (2.00–2.10) �n pre-vent length; he�ght 0.77 (0.74–0.79) t�mes w�dth. Snout very long, narrowly tr�angular �n lateral v�ew, acutely angular �n dorsal v�ew, hor�zontal length 1.85 (1.61–1.84) t�mes eye length, 0.98 (0.95–1.01) t�mes �nterorb�tal space; hor�zontal prenar�al length 1.85 (1.88–2.01) t�mes �n preoral length; apex narrowly rounded. Eye broadly oval, relat�vely large, length 5.17 (4.77–5.44) �n head, 1.53 (2.13–2.63) t�mes he�ght. Sp�racle small, horseshoe-shaped; broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.23 (2.68–3.89) �n eye length. G�ll open�ngs sl�ghtly obl�que d�rected sl�ghtly posteroventrally, subequal in size, height of fifth slit 1.98 (1.7–2.2)% TL. Mouth almost transverse, upper jaw strongly arched, �ts w�dth 1.92 (1.72–1.91) �n preoral length; three ser�es of funct�onal teeth �n upper jaw, two ser�es �n lower; upper lab�al furrows about tw�ce length of lower furrows, prom�nent groove extend�ng poster�or-laterally from angle of jaws. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Nostr�ls small, almost transverse; anterior nasal flap bifurcate, upper lobe largest; posterior lobe narrow, sk�rt-l�ke; �nternar�al space 2.56 (2.33–2.82) �n preoral length, 2.61 (2.20–3.14) t�mes nostr�l length. Dermal dent�cles on back very small, weakly tr�cusp�date w�th pronounced med�an r�dge; lateral cusps vary�ng from barely detectable to short, weakly angular; on dorsal and caudal fins leaf-shaped, strongly imbricated, more regular on snout t�p; �nterorb�t pr�mar�ly w�th un�cusp�date dent�cles, sharply po�nted poster�orly, w�th three broad lobe-l�ke expans�ons on lateral and anter�or marg�n, med�an r�dge pronounced; on belly very strongly �mbr�cated, pungent d�stally, un�cusp�date to weakly tricuspidate, ridges appear less pronounced; dorsal fins ent�rely covered w�th closely spaced dent�cles; dent�cles present over g�ll membranes; dent�cles around sp�racle un�cusp�date, extend�ng �nto cav�ty anter�orly, fold covered with denticles. First dorsal fin low, raked slightly, weakly rounded ap�cally; anter�or marg�n moderately convex; poster�or marg�n moderately concave, upper port�on d�rected sl�ghtly anterodorsally; free rear t�p relat�vely th�ck, short; �nner marg�n almost stra�ght; �nsert�on of base well forward of pelvic-fin origin; fin-spine origin above pectoral-fin rear tip; spine base moderately broad, exposed anter�orly well below junct�on of sp�ne and fin; spine tapering rapidly distally, smaller than second


A

B

Figure 1. Lateral v�ew of: A. Squalus nasutus sp. nov. holotype (CSIRO H 2590–12, female 508 mm TL); B. Squalus japonicus (CSIRO H 6294–31, female 571 mm TL).

dorsal-fin spine; pre-first dorsal length 3.06 (3.03–3.22) times in TL; first dorsal-fin length 2.11 (1.89–2.11) t�mes �ts he�ght, 1.25 (1.18–1.23) t�mes second dorsal-fin length; first dorsal-fin height 1.77 (1.47–1.74) times second dorsal-fin height; exposed first dorsal spine length 0.48 (0.36–0.54) times height of fin. Second dorsal fin small, strongly raked; poster�or marg�n deeply concave, upper port�on d�rected sl�ghtly dorsoposter�orly; free rear tip long, inner margin length 1.43 (1.01–1.34) times fin height; second dorsal-fin length 2.98 (2.53–3.00) times its height; spine length 1.07 (0.97–1.17) in height of fin; fin-spine origin well behind insertion of pelvic fin, exposed at level of junction with spine and fin; interdorsal space 1.02 (0.92–0.99) in length from snout tip to pectoral-fin origin, 1.31 (1.21–1.36) in pre-first dorsal length; second sp�ne w�th very broad base, slender d�stally, taper�ng very rapidly just above point of exposure. Pectoral fin small, anter�or marg�n weakly convex; �nner marg�n moderately convex, length 7.1 (6.6–7.7)% TL; apex narrowly rounded, lobe-l�ke but not falcate; poster�or marg�n moderately convex, free rear t�p bluntly angular; base very short, 2.74 (2.85–3.11) in length of anterior margin. Pelvic fins small, anter�or and poster�or marg�ns almost stra�ght, apex broadly rounded, free rear t�p narrow, acute. Caudal peduncle long, tapering slightly to caudal fin, pelvic–caudal space 0.88 (0.70–0.82) �n pectoral–pelv�c space, 1.03 (0.87–0.96) �n prepectoral length; dorsal–caudal space 2.26 (2.06–2.47) �n �nterdorsal length; lateral folds extending from below insertion of second dorsal fin to about a pupil diameter posterior of caudal pits. Caudal fin relat�vely short, dorsal caudal marg�n 1.28 (1.14–1.26) �n head length; length of lower caudal lobe 1.60 (1.59–1.79)

�n upper lobe length. Vertebral centra 107 (103–109 �n 18 paratypes), monospondylous 39 (36–39), precaudal 81 (78–83) and caudal 26 (23–28). Teeth �n upper jaw (of paratype, CSIRO H 2598–07) 13+13=26, lower jaw 11+11=22.

COLOUR.— When fresh (based on holotype): dorsal surface grey�sh brown, much paler ventrally; l�ght and dark tonal areas strongly demarcated on head, the�r border extend�ng from snout, beneath eye, through top of g�ll slits to pectoral-fin origin; tonal areas poorly demarcated on trunk. Dorsal fins uniformly greyish brown, anterior bases and free rear t�p sl�ghtly paler; sp�nes dusky, anter�or marg�ns sl�ghtly darker; sk�n at base of exposed spine black. Caudal fin mostly dusky; black caudal bar subvert�cal, prom�nent, broad, extend�ng from fork to about halfway along poster�or marg�n of upper lobe; upper halves of dorsal and ventral lobes off-wh�te; black blotches at base of upper and lower lobes. Pectoral fin greyish dorsally, with well-defined white posterior margin; pelvic fins similar, ventral surface pale grey with a wh�te base and poster�or marg�n. Juven�le paratype (CSIRO H 1652–01) s�m�lar to holotype. but w�th much more pronounced fin markings; dorsal fins with black outer halves. Caudal bar relat�vely short, broad, black, slightly taller than eye length; upper caudal-fin lobe with well-defined black blotch on anterior margin and weak upper med�an dusky blotch; ventral lobe paler than apex of upper lobe. In preservat�ve: holotype s�m�lar, but l�ght and dark tonal areas on body less d�st�nct; dark mark�ngs on fins faint but evident; broad, pale posterior borders on pectoral and pelvic fins well-defined.

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Figure 2. Ventral v�ew of the head of A. Squalus nasutus sp. nov. holotype (CSIRO H 2590–12, female 508 mm TL); B. Squalus japonicus (HUMZ 80224, female 641 mm TL).

SIZE.— Austral�an females and males atta�n at least 590 (non-type) and 499 mm TL (non-type), respect�vely; smallest adult male 461 mm TL, st�ll adolescent at 452 mm TL. Ten adult males from Indones�a and the Ph�l�pp�nes ranged from 464–539 mm TL; 25 females from 470–634 mm TL.

DISTRIBUTION.— Upper cont�nental slope off Western Austral�a, north of the Rottnest Canyon (ca 32°S, 115°E) to the Rowley Shoals (ca 17°S, 120°E). Occurs �n depths of 300–850 m but trawled ma�nly �n depths of 300–400 m. Appears to be d�str�buted through the Indones�an Arch�pelago to the Ph�l�pp�nes but depth

A

B


Table 1. Proport�onal d�mens�ons as percentages of total length for the holotype (CSIRO H 2590–12) and 5 paratypes of Squalus nasutus sp. nov., w�th compar�sons w�th Indones�an and Ph�l�pp�nes mater�al.

Western Australia Indonesia Philippines Holotype Paratypes n = 4 n = 3

M�n. Max. M�n. Max. M�n. Max.TL – Total length 508 452 470 468 576 473 541PCL – Precaudal length 81.7 79.4 80.9 80.6 81.0 80.4 81.0PD2 – Pre-second dorsal length 65.4 63.1 64.4 64.5 66.3 63.4 64.1PD1 – Pre-first dorsal length 32.7 31.0 33.0 31.7 34.4 31.8 33.5SVL – Pre-vent length 51.8 46.9 50.5 48.4 49.3 47.8 49.7PP2 – Prepelv�c length 48.8 45.3 47.7 45.5 47.9 45.5 48.4PP1 – Prepectoral length 25.6 23.3 24.2 23.7 24.7 24.6 25.2HDL – Head length 23.6 23.0 24.1 23.2 24.1 23.5 24.5PG1 – Prebranch�al length 21.0 19.4 20.8 19.2 20.5 19.6 20.7PSP – Presp�racular length 14.9 13.4 14.8 13.3 14.7 14.0 14.9POB – Preorb�tal length 9.9 8.5 9.3 8.3 9.1 9.3 10.1PRN – Prenar�al length 7.3 6.4 6.7 5.9 6.6 6.8 7.5POR – Preoral length 12.7 11.4 12.3 11.1 12.0 11.7 12.4INLF – Inner nostr�l-lab�al furrow space 4.9 4.8 5.2 4.7 5.1 4.6 5.1MOW – Mouth w�dth 6.6 6.3 7.2 6.0 6.6 5.8 6.2ULA – Lab�al furrow length 2.3 1.8 2.2 1.9 2.3 1.8 2.2INW – Internar�al space 5.0 4.2 5.1 4.3 4.5 4.3 4.7INO – Interorb�tal space 8.6 7.8 8.6 7.3 7.5 7.3 7.9EYL – Eye length 4.6 4.3 5.1 4.5 5.0 4.5 4.9EYH – Eye he�ght 3.0 1.9 2.1 1.8 2.2 1.9 2.1SPL – Sp�racle length 1.4 1.3 1.6 1.3 1.7 1.5 1.7GS1 – F�rst g�ll-sl�t he�ght 1.7 1.3 1.6 1.4 1.6 1.1 1.6GS5 – F�fth g�ll-sl�t he�ght 2.0 1.7 2.2 2.0 2.2 1.7 1.9IDS – Interdorsal space 25.0 24.4 25.6 25.8 28.0 25.6 26.1DCS – Dorsal-caudal space 11.1 10.4 12.0 9.8 10.8 10.4 11.5PPS – Pectoral-pelv�c space 21.9 18.7 20.8 18.4 22.5 17.8 22.0PCA – Pelv�c-caudal space 24.9 25.2 26.6 25.2 27.7 25.4 26.2D1L – F�rst dorsal length 12.5 12.0 12.8 10.6 11.6 11.1 11.8D1A – F�rst dorsal anter�or marg�n 10.6 10.4 10.6 8.9 10.0 9.3 10.1D1B – F�rst dorsal base length 7.1 6.8 7.7 5.6 6.9 6.4 6.9D1H – F�rst dorsal he�ght 5.9 5.7 6.8 5.9 6.2 6.2 6.5D1I – F�rst dorsal �nner marg�n 5.6 4.9 5.5 4.7 5.3 4.5 5.3D1P – F�rst dorsal poster�or marg�n 7.2 6.6 8.4 6.8 7.4 7.5 7.7D1ES – F�rst dorsal sp�ne length 2.8 2.2 3.4 3.8 3.8 3.2 3.4D1BS – F�rst dorsal sp�ne base w�dth 0.9 0.7 1.0 0.6 0.9 0.7 0.8D2L – Second dorsal length 10.0 9.9 10.9 9.3 9.9 9.5 10.3D2A – Second dorsal anter�or marg�n 8.3 7.5 8.9 7.1 8.3 8.0 8.9D2B – Second dorsal base length 5.4 5.2 6.6 4.9 6.0 5.9 6.4D2H – Second dorsal he�ght 3.3 3.3 4.2 3.3 3.8 3.7 4.2D2I – Second dorsal �nner marg�n 4.8 3.9 5.0 3.8 4.5 3.4 4.4D2P – Second dorsal poster�or marg�n 4.6 4.4 5.3 3.9 4.8 4.3 4.9D2ES – Second dorsal sp�ne length 3.6 3.5 4.7 3.2 3.9 4.0 4.5D2BS – Second dorsal sp�ne base w�dth 1.1 0.9 1.1 0.7 0.9 0.6 0.9

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Table 1. cont’d.

Western Australia Indonesia Philippines Holotype Paratypes n = 4 n = 3

M�n. Max. M�n. Max. M�n. Max.P1A – Pectoral anter�or marg�n 12.9 12.0 13.3 11.4 13.9 12.0 13.0P1I – Pectoral �nner marg�n 7.1 6.6 7.7 5.9 6.4 5.9 6.8P1B – Pectoral base length 4.7 4.0 4.3 3.9 4.9 4.5 4.8P1P – Pectoral poster�or marg�n 8.2 7.0 8.5 8.4 10.4 7.7 8.7P2L – Pelv�c length 10.2 9.0 12.1 9.8 12.0 8.3 11.2P2H – Pelv�c he�ght 4.3 4.0 4.9 3.9 6.0 4.2 4.4P2I – Pelv�c �nner marg�n 4.7 3.8 6.6 4.6 6.6 3.8 7.3CDM – Dorsal caudal marg�n 18.5 19.0 20.1 18.8 19.2 19.4 19.7CPV – Preventral caudal marg�n 11.6 10.6 12.3 10.6 11.2 10.7 10.9CPU – Upper postventral caudal marg�n 13.2 13.2 14.3 13.5 13.8 13.6 14.4CPL – Lower postventral caudal marg�n 4.7 3.8 4.5 4.2 5.1 4.1 5.1CFW – Caudal fork w�dth 6.7 6.3 6.7 6.2 6.5 6.3 6.4CFL – Caudal fork length 9.3 8.5 9.7 8.0 8.8 7.9 9.2HANW – Head w�dth at nostr�ls 8.3 8.1 8.5 7.6 8.0 7.8 8.0HAMW – Head w�dth at mouth 11.0 10.7 11.5 9.8 10.9 9.8 10.3HDW – Head w�dth 12.6 11.4 11.9 11.5 12.4 11.5 11.9TRW – Trunk w�dth 11.0 8.6 9.6 8.9 10.6 8.3 9.2ABW – Abdomen w�dth 9.7 7.9 9.6 8.3 9.9 6.9 8.7TAW – Ta�l w�dth 6.9 5.9 6.6 5.0 6.9 6.0 6.5CPW – Caudal peduncle w�dth 3.0 2.9 3.5 2.8 3.0 2.9 3.1HDH – Head he�ght 9.6 8.7 9.1 8.2 9.2 8.7 9.7TRH – Trunk he�ght 11.4 9.4 10.2 9.2 10.5 10.1 11.2ABH – Abdomen he�ght 12.2 8.6 10.8 7.8 10.9 10.4 10.6TAH – Ta�l he�ght 7.0 6.0 7.7 5.4 6.7 6.1 7.5CPH – Caudal peduncle he�ght 2.2 2.2 2.3 2.0 2.1 2.3 2.6CLO – Clasper outer length – 2.5 4.5 4.4 4.4 – 4.6CLI – Clasper �nner length – 9.3 12.2 11.4 11.4 – 11.4CLB – Clasper base w�dth – 1.3 1.6 1.5 1.5 – 1.5

d�str�but�on unknown as exam�ned mater�al was collected from fish markets.

ETYMOLOGY.— Der�ved from the Lat�n nasutus mean�ng ‘large-nosed’ �n allus�on to �ts elongated prenar�al snout.

VERNACULAR.— Western Longnose Spurdog.

REMARKS.— The type ser�es of Squalus nasutus �s based on Austral�an mater�al because m�nor d�fferences �n morphometr�cs (Table 1) and mer�st�cs ex�st between populations across the Indo–West Pacific. Apart from hav�ng sl�ghtly h�gher total vertebral counts �n Ph�l�pp�ne spec�mens (109–111, n=3 vs. 103–109 �n Austral�an forms, n=10), the Indo–Ph�l�pp�ne spec�mens appear to have a sl�ghtly longer �nterdorsal space (25.6–28.0 vs.

Figure 3. Cusps of the flank denticles of Squalus nasutus sp. nov. paratype (CSIRO H 1207–08, female 496 mm TL). F�eld of v�ew w�dth 1.2 mm.


Figure 4. Lateral view of the dorsal fins of: Squalus nasutus sp. nov. paratype (CSIRO H 2598–07, female 465 mm TL) – A. first dorsal fin, B. second dorsal fin.

A

B

24.4–25.6% TL), sl�ghtly narrower �nterorb�t (7.3–7.9 vs. 7.8–8.6% TL) and head (w�dth at nostr�ls 7.6–8.0 vs. 8.1–8.5% TL), and a marginally shorter pectoral-fin inner margin (5.9–6.8 vs. 6.6–7.7% TL) and first dorsal fin (length 10.6–11.8 vs. 11.1–12.8% TL). More mater�al �s needed to understand the extent of �nterreg�onal var�at�on between populat�ons.

The new spec�es belongs to a subgroup of Squalus that �ncludes 3 other, long-snouted spec�es: S. japonicus (western North Pacific), S. melanurus (New Caledon�a) and S. rancureli (New Caledon�a). Squalus nasutus �s much smaller than these spec�es, atta�n�ng only 63 cm TL w�th males matur�ng at about 45 cm TL, whereas the other spec�es are reported to reach at least 91 cm, 75 cm, and 77 cm TL, respect�vely, and have females that mature larger than 65 cm TL (Compagno, 1984). Of these spec�es, Squalus nasutus appears closest to S. japonicus (F�gs 1b, 2b, 5b) from the western North Pacific with which it appears to have been confused in the Ph�l�pp�nes (Compagno et al., 2005b). These spec�es differ significantly from the two New Caledonian endem�cs: from S. melanurus �n lack�ng prom�nent dark black markings on the dorsal and caudal fins; and from S. rancureli �n hav�ng a more acutely po�nted snout, a longer ventral caudal lobe and possess�ng a well-developed secondary lobe on the anterior nasal flap.

Squalus nasutus d�ffers from S. japonicus �n both morphometr�cs and mer�st�cs. Chen et al. (1979) g�ves

A

B

Figure 5. Juvenile coloration of the caudal fin of: A. Squalus nasutus sp. nov. (CSIRO H 1693–02, female 306 mm TL); B. Squalus japonicus (UF 148932, male embryo 247 mm TL).

relat�vely h�gher vertebral counts for S. japonicus (40–45 vs. 36–39 monospondylous centra for S. nasutus from Austral�a; 82–91 vs. 78–81 precaudal centra; and 110–119 vs. 103–109 total centra). There also appears to be a d�fference �n the rat�o of d�plospondylous trunk to monospondylous centra (0.8 based on one of Chen et al.’s spec�mens vs. 1.1–1.2 �n the 10 types of S. nasutus). Teeth counts appear s�m�lar, �n one paratype w�th 26 �n the upper jaw (vs. 25–27, ma�nly 27 �n 33 spec�mens) and 22 �n the lower jaw (vs. 23–24, ma�nly 23 �n 33 spec�mens). The most obv�ous morphometr�c d�fferences ex�st around the head, pectoral fin and trunk. The pectoral fin of adult S. japonicus �s much larger, much more falcate w�th a more concave poster�or marg�n (F�g 2b), and a longer �nner marg�n (8.6–11.1 vs. 6.6–7.1% TL �n S. nasutus). The preoral length (10.2–11.4 vs. 11.1–11.7% TL) and eye (length 3.8–4.2 vs. 4.3–5.1% TL) of S. japonicus are relat�vely smaller, and the trunk appears to be more depressed (trunk he�ght 8.2–9.5 vs. 9.4–11.4% TL; trunk w�dth 12.0–12.6 vs. 8.6–11.0% TL �n S. nasutus). Key

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measurements taken by Ish�kawa (1908) of the male holotype of S. japonicus (700 mm TL) conformed well to the ranges prov�ded by Chen et al. (1979) for th�s spec�es. He recorded a preoral length of 75 mm (10.7% TL) and a 26 mm eye (3.7% TL), both of wh�ch conform to the ranges l�sted above.

Other material.Squalus japonicus: HUMZ 39455, female 535 mm TL, HUMZ 48406, female 339 mm TL, HUMZ 80501, �mmature male 291 mm TL, HUMZ 97257, adult male 602 mm TL, Mimase fish market, Kochi, Japan; HUMZ 80224, female 641 mm TL, Ok�nawa Trough, Japan, 310 m; CSIRO H 6294–26, male 456 mm TL, CSIRO H 6294–27, female 380 mm TL, CSIRO H 6294–31, female 571 mm TL, UF 148932, male embryo 247 mm TL, Tashi fish market, north-east Taiwan.

ACKNOWLEDGEMENTS

Th�s study was conducted �n parallel w�th a project on southern Australian dogfishes funded by the Fisheries Research and Development Corporat�on (FRDC 1998/108) and the new species was first identified in a study of the Austral�an chondr�chthyan fauna (Last and Stevens, 1994), also funded by the same agency (FRDC). S�ncere thanks go to Alasta�r Graham for assembl�ng data on Austral�an mater�als, Lou�se Conboy for ed�t�ng �mages of the types and other mater�al, Matt Edmunds for h�s extens�ve research on th�s group, Tony Rees for ass�stance w�th preparat�on of SEMs, T�m Founta�n for prepar�ng rad�ographs and obta�ned mer�st�c �nformat�on, and Gordon Yearsley, John Stevens and John Pogonosk� for the�r ed�tor�al comments. We thank curators and curator�al staff for ass�stance w�th obta�n�ng spec�mens and collect�on data: Mart�n Gomon and D� Bray (NMV), Barry Hutch�ns, Sue Morr�son and Glenn Moore (WAM), Mark McGrouther (AMS), ‘May’ Luchavez-Maypa, ‘Adon’ Guad�ano (SUML), George Burgess (UF) and Kazu Nakaya and H�sash� Imamura (HUMZ). Indones�an mater�al was collected as part of a 5-year ACIAR (Austral�an Centre for Internat�onal Agr�cultural Research) funded project and �mportant contr�butors to th�s project �nclude, Junaed� (Kedonganan F�sh Market, Bali), processors and fishers at Tanjung Luar fish market (Lombok), Fahm� (LIPI), Dharmad� (RCCF), Jenny G�les (QDPI&F), Ono Kurnaen Sumadh�harga (LIPI), Subhat Nurhak�m (RCCF), Ian Potter (Murdoch Un�vers�ty), Cathy D�chmont and Steve Blaber (CSIRO). Ph�l�pp�nes mater�al was collected as part of a WWF funded project and �mportant contr�butors to th�s project �nclude, Andy Ol�ver (formerly WWF, USA) and Moonyeen Alava (formerly WWF Ph�l�pp�nes).

REFERENCES

Chen, C., Tan�uch�, T. and Nose, Y. (1979) Bla�nv�lle’s dogfish, Squalus blainville, from Japan, w�th notes on S. mitsukurii and S. japonicus. Japanese Journal of Ichthyology, 26(1), 26–42.


Compagno, L.J.V., Dando, M. and Fowler, S. (2005a) A Field Guide to the Sharks of the World. Harper Coll�ns Publ�sh�ng Ltd., London, 368 pp.

Compagno, L.J.V., Last, P.R., Stevens, J.D. and Alava, M.N.R. (2005b) Checkl�st of Ph�l�pp�ne Chondr�chthyes. CSIRO Marine Laboratories Report 243, 103 pp.

Fourmano�r, P. and I. R�vaton, I. (1979) Po�ssons de la pente réc�fale externe de Nouvelle-Calédon�e et des Nouvelles-Hébr�des. Cahiers de l’Indo-Pacifique, 1(4), 405–443.

Ish�kawa, C. (1908) Descr�pt�on of a new spec�es of squalo�d shark from Japan. Proceedings of the Academy of Natural Sciences of Philadelphia, 60, 71–73.

Last, P.R. and Stevens, J.D. 1994. Sharks and Rays of Australia. CSIRO, Austral�a, 513 pp.

Last, P.R., Wh�te, W.T. and Stevens, J.D. (2007) Part 5 — New spec�es of Squalus of the ‘highfin megalops group’ from the Australas�an reg�on, p. 39–53. In: Descr�pt�ons of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last P.R., Wh�te W.T. and Pogonosk� J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.




Part 9 — Redescription of the Northern Spiny Dogfish Squalus griffini Phillipps, 1931 from New Zealand

Cl�nton A. J. Duffy1 and Peter R. Last2

1Department of Conservat�on, Pr�vate Bag 68908, Newton, Auckland, 1145, NEW ZEALAND2CSIRO Mar�ne & Atmospher�c Research, GPO Box 1538, Hobart, TAS, 7001, AUSTRALIA

ABSTRACT.— The Northern Spiny Dogfish, Squalus griffini Ph�ll�pps, from central and northern New Zealand �s resurrected based upon deta�led re-exam�nat�on of the holotype and new mater�al collected from throughout �ts range. Compar�sons are made w�th S. griffini and other large Indo–Pacific species of Squalus, �nclud�ng S. mitsukurii from the western North Pacific, and newly recognised members of the ‘mitsukurii group’ from Austral�a. Squalus griffini d�ffers from members of the ‘m�tsukur�� group’ �n hav�ng a wh�te posterior caudal-fin margin without a dark caudal bar. It also appears to be atypically variable in body and fin shapes, adult coloration and denticle morphology.

Key words. Squalo�dea – Squal�dae – Squalus griffini – resurrected spec�es – New Zealand – Australas�a


INTRODUCTION

Two spec�es of Squalus are recogn�sed from New Zealand waters. Squalus acanthias (L�nnaeus, 1758), known as the Spotted Spiny Dogfish (New Zealand) or Whitespotted Dogfish (Australia), is widely distributed around North and South Islands of New Zealand, as well as over the Chatham R�se and Campbell Plateau, where �t �s most abundant south of 40° S (Anderson et al., 1998). The Northern Spiny Dogfish, presently referred to as S. mitsukurii Jordan and Snyder �n Jordan and Fowler, 1903 (Paul�n et al., 1989), �s restr�cted to waters north of the Subtrop�cal Front. It occurs along the cont�nental marg�n off the west coasts of the North and South Islands from about 34° S to 44° S, w�th some �solated records from research trawls on the central Challenger Plateau. Off the east coast, �t has been recorded southward to almost 45° S but �s rarely caught south of 38° S, except around the Chatham Islands between 175° and 179° W (Anderson et al., 1998).

The Northern Spiny Dogfish was originally recorded erroneously from the Northland as S. megalops by Regan (1914) but was later descr�bed as a new spec�es, S. griffini, by Ph�ll�pps (1931). Ph�ll�pps’ descr�pt�on, based upon two spec�mens from the Haurak� Gulf, was meager and unaccompan�ed by an �llustrat�on. He noted that a descr�pt�on of S. griffini and two new Austral�an spec�es was be�ng publ�shed elsewhere and s�mply descr�bed �t as show�ng no s�gn of spots and hav�ng a relat�vely mass�ve compressed spine in front of the second dorsal fin. He d�st�ngu�shed S. griffini from S. megalops pr�mar�ly by a heav�er, more compressed and unpol�shed appearance of

the second dorsal sp�ne, a somewhat shorter head, and a more posterior location of the pelvic fin. Whitley (1940) prov�ded a sl�ghtly more deta�led descr�pt�on of S. griffini, report�ng three proport�onal measurements and repeat�ng Ph�ll�pps’ (1931) descr�pt�on of the second dorsal sp�ne and morpholog�cal compar�son w�th S. megalops. He also figured the species for the first time, reproducing a copy of a pa�nt�ng of S. griffini prov�ded by Ph�ll�pps. Ph�ll�pps’ (1946) account of the species figured the original painting of the type spec�men by E. H. Atk�nson but added l�ttle to the descr�pt�on, not�ng only that the centre of the pelv�c-fin base is midway between the posterior orbit and the caudal-fin tip (rather than midway between the snout and caudal t�p as �n S. megalops).

Garr�ck (1960) rev�ewed the Australas�an spec�es of Squalus and �ncorrectly synonym�sed S. griffini and S. fernandinus Mol�na, 1782 (sensu B�gelow and Schroeder, 1948, 1957) w�th S. blainvillii (�ncorrect spell�ng of S. blainville (R�sso, 1827), Eschmeyer, 1998). Garr�ck recogn�sed S. acanthias and S. blainville from New Zealand, and S. acanthias, S. blainville and S. megalops from Austral�a. Squalus blainville was thought to be w�despread �n the Atlant�c, Ind�an and Pacific Oceans (Bigelow and Schroeder, 1948, 1957; Garr�ck 1960). However, �n a rev�ew of Japanese Squalus, Chen et al. (1979) defined S. blainville as a spec�es w�th high dorsal fins and long dorsal-fin spines based upon exam�nat�on of Japanese mater�al and descr�pt�ons of S. blainville purportedly from �ts type local�ty, the Med�terranean. They observed that Squalus, referred to S. fernandinus and S. blainville by B�gelow and Schroeder (1948) and Garrick (1960), had short dorsal-fin spines

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and were more s�m�lar to S. mitsukurii from Japan, and suggested that nom�nal S. blainville from New Zealand could be �dent�cal to S. mitsukurii. Compagno (1984) also noted that dogfishes resembling S. mitsukurii occurred off Austral�a and New Zealand, and d�d not recogn�se S. blainville from the Southern Hem�sphere. Although none of the above authors made d�rect compar�sons between S. mitsukurii and New Zealand mater�al, the Northern Spiny Dogfish has subsequently been widely synonym�sed w�th S. mitsukurii (Paul�n et al. 1989; Yano �n Amaoka et al., 1990; Roberts, 1991; Paul and Heath, 1997; Cox and Franc�s, 1997; Anderson et al., 1998; Compagno et al., 2005). Last and Stevens (1994) prov�s�onally mapped New Zealand populat�ons as S. mitsukurii but suggested that reg�onal forms may represent a spec�es complex.

We exam�ned the holotype of S. griffini and add�t�onal mater�al held �n the New Zealand and Austral�an Nat�onal Fish Collections, and concluded that these are specifically d�st�nct from Austral�an mater�al once �ncorrectly referred to as S. mitsukurii (see Part 6 of th�s �ssue, Last et al., 2007a), as well as mater�al of S. mitsukurii from the western North Pacific. In this paper, S. griffini �s redescr�bed based upon a reexam�nat�on of the holotype and add�t�onal mater�al collected from the Lou�sv�lle R�dge, Kermadec R�dge, Wanganella Bank, Chatham R�se, and North and South Islands of New Zealand.

METHODS

Morphometr�c characters were selected to enable compar�sons to be made w�th other Squalus taxa treated �n the ser�es of papers �n th�s �ssue (Part 1 of th�s �ssue, Last et al., 2007b). The holotype and 16 spec�mens from the North and South Islands of New Zealand, the Wanganella Bank, Chatham R�se, and southern Kermadec and Lou�sv�lle R�dges were measured �n full (Table 1). Vertebral counts were taken from X-rays of 10 spec�mens (�nclud�ng the holotype). Tooth row counts were taken d�rectly from spec�mens. Morphometr�c and mer�st�c values for the holotype are g�ven, followed by ranges for other spec�mens �n parentheses. The holotype and a large collect�on of spec�mens are held at the Nat�onal F�sh Collect�on, Museum of New Zealand, Te Papa Tongarewa, Well�ngton (NMNZ); 3 add�t�onal spec�mens used �n th�s study are held at the Austral�an Nat�onal F�sh Collect�on, Hobart (CSIRO). Spec�men reg�strat�on numbers are prefixed by these acronyms.

Squalus griffini Ph�ll�pps, 1931


Squalus megalops: (not Macleay) Regan, 1914: 14; Ph�ll�pps, 1927: 9.Squalus griffini Phillipps, 1931: 360; Phillipps, 1946: 16, fig. 5;

Moreland, 1957; Parrott, 1958: 114, �ncorrectly �llustrated w�th a figure of S. megalops from McCulloch (1922).Flakeus griffini: Whitley, 1940: 136, fig. 149; Powell, 1951: 61, fig. 295.Squalus blainvillii: (not Risso) Garrick, 1960: 532–537, figs 1D–F, 3A–F, 6A–F.Squalus blainville: (not R�sso) Garr�ck, 1961; Doogue et al., 1966; York, 1970; Paul, 1985: 21, fig. 7; Ayling and Cox, 1982: 65, fig. Squalus mitsukurii: (not Jordan and Snyder) Compagno, 1984: 121–122, figs; Paulin et al., 1989: 13, fig. 7.3; Roberts, 1991: 8, fig. 4; Last and Stevens, 1994: 101–102, 8.39, Plate 5; Paul and Heath, 1997: 2, fig.; Cox and Francis, 1997: 45, fig.

Holotype. NMNZ P 662, adult female 972 mm TL, Haurak� Gulf, North Island, New Zealand, 1931.Other material. 20 spec�mens. NMNZ P 2646, female 1025 mm TL, off Foxton, New Zealand, 40°28′ S, 175°14′ E, 55 m; NMNZ P 2649, female 955 mm TL, off Kapiti Island, New Zealand, 40°51′ S, 174°52′ E, 37 m; NMNZ P 2759, �mmature female 539 mm TL, off Lott�n Point, 37°32′ S, 178°10′ E, New Zealand, 137 m; NMNZ P 5176, female 929 mm TL, near Cuv�er Island, New Zealand, 36°26′ S, 175°46′ E, 77 m; NMNZ P 20965, female 482 mm TL, east of the Chatham Islands, New Zealand, 43°20′ S, 176°36′ W, 136 m; NMNZ P 22549, adult male 794 mm TL, Lou�sv�lle R�dge, New Zealand, 32°00′ S, 172°00′ W; NMNZ P 35270, adult male 867 mm TL, Chatham Rise, New Zealand, 43°43′ S, 175°22′ W, 543 m; NMNZ P 35973, female 823 mm TL, southern Kermadec Ridge, New Zealand, 32°43′ S, 179°20′ W, 333 m; NMNZ P 39893, female 1018 mm TL, NMNZ P 39899, near-term male embryo 227 mm TL, off Volkner Rocks, Wh�te Island, Bay of Plenty, New Zealand, 37°28′ S, 177°08′ E, 265 m; NMNZ P 40888, adult female 1070 mm TL, off Nap�er Island, Kermadec Islands, New Zealand, 29°14′ S, 177°53′ E, 350 m; NMNZ P 41774, adult male 693 mm TL and 4 females 737–835 mm TL, Southern Challenger Plateau, New Zealand, 41°28′ S, 171°06′ E, 175m; NMNZ P 41775, �mmature female 418 mm TL, Wa�rarapa, New Zealand, 40°15′ S, 177°01′ E, 616 m; CSIRO H 6057–12, female 836 mm TL, Tasman Sea, West Norfolk R�dge, 33° 45′ S, 167°17′ E, 259 m; CSIRO H 5002–06, adult male 819 mm TL, CSIRO H 5002–07, female 974 mm TL, Wanganella Bank, Norfolk Ridge, 31°49′ S, 167°47′ E, 351 m; CSIRO H 6006–11, �mmature male 473 mm TL, Norfolk Ridge, south of Norfolk Island, 29°42′ S, 168° 02′ E, 344 m; CSIRO H 6066-08, immature male 262 mm TL, Wanganella Bank, Norfolk Ridge, 32°33′ S, 167°38′ E, 362 m; CSIRO H 6069–02, adult male 789 mm TL, Wanganella Bank, Norfolk Ridge 32°35′ S, 167°41′ E, 497 m; NMV A 25109–007, �mmature male 531 mm TL, mid-Norfolk Ridge 29°42′ S, 168°02′ E, 344 m.

DIAGNOSIS.— A large spec�es of Squalus w�th the follow�ng comb�nat�on of characters: body slender, depth 8.6–13.9% TL; snout long, preoral length greater than mouth w�dth, 2.4–2.6 t�mes hor�zontal prenar�al length, 8.8–11.4% TL; eye large, 3.6–5.3% TL; secondary


A

B

C

Figure 1. Lateral v�ew of: A. Squalus griffini (NMNZ P 39893, female 1018 mm TL); B. Squalus griffini holotype (NMNZ P 662, female 972 mm TL); C. Squalus griffini (CSIRO H 6006–11, �mmature male 473 mm TL).

lobe of anterior nasal flap well developed; first dorsal fin moderate-sized, height 6.3–8.7% TL, triangular, poster�or marg�n almost stra�ght, sp�ne short and heavy; second dorsal fin raked, posterior margin deeply concave, he�ght up to 4.2–5.8% TL, sp�ne long w�th strong base; pectoral fin of adult not falcate; pectoral-fin inner margin short, 5.4–7.7% TL; caudal fin with broad pale posterior marg�n and lower lobe �n all but largest adults, dark caudal bar absent, well-defined stripes above and below soft portion of fin in juveniles; flank denticles tricuspid; 45–47 monospondylous centra, 86–91 precaudal centra, 113–121 total centra; adult max�mum s�ze at least 110 cm TL.

DESCRIPTION.— Body elongate fus�form to sl�ghtly compressed; head long 21.0 (20.2–24.3)% TL; caudal peduncle elongate, narrow, depressed sl�ghtly, 23.8 (21.7–26.0)% TL. Head depressed forward of sp�racles, length 2.48 (2.07–2.51) �n pre-vent length; he�ght 0.76 (0.60–0.82) w�dth. Snout long, tr�angular �n lateral v�ew,

narrowly rounded �n dorsal v�ew, hor�zontal length 1.87 (1.16–1.92) t�mes eye length, 0.94 (0.67–0.88) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.42 (2.37–2.65) �n preoral length; apex narrowly rounded. Eye broadly oval, relat�vely large, length 5.88 (4.29–5.91) �n head, 2.60 (1.83–3.46) t�mes he�ght. Sp�racle small, broadly crescent�c; broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 2.43 (2.25–3.61) �n eye length. G�ll open�ngs small, obl�que, d�rected sl�ghtly posteroventrally; subequal in size, height of first gill slit 2.06 (1.73–2.50)% TL; fifth gill slit wrapping around pectoral fin origin. Nostrils small, almost transverse; anterior nasal flap strongly bifurcate, weakly bifurcate �n some other mater�al; poster�or lobe narrow, sk�rt-l�ke; �nternar�al space 2.09 (1.97–2.31) t�mes �n preoral length, 3.03 (2.48–3.22) t�mes nostr�l length. Mouth arched, w�dth 1.62 (1.43–1.88) �n preoral length; upper lab�al furrows 2.2 (1.8–2.7)% TL, cont�nued as prom�nent grooves extend�ng poster�or-laterally from angle of jaws; lower furrows about half the length of the uppers, generally not

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A

B

Figure 2. Ventral v�ew of the head of: A. Squalus griffini holotype (NMNZ P 662, female 972 mm TL); B. Squalus griffini (CSIRO H 6006–11, �mmature male 473 mm TL).

v�s�ble �n ventral v�ew. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke; cusps d�rected strongly laterally, low, base of tooth broader than length of cusp. Dermal denticles on flank below first dorsal fin of holotype broad, strongly imbricated, tr�cusp�d, w�th weak lateral cusps and well-developed, grooved lateral r�dges; var�able �n other mater�al, dent�cles often weakly �mbr�cate w�th long cusps w�th weak lateral r�dges, even �n adult males and large females of s�m�lar s�ze to the holotype; present on g�ll membranes. F�rst dorsal fin of moderate size, broadly rounded apically, poster�or marg�n almost upr�ght, strongly convex near fin base; free rear tip base relatively thick, short; inner margin almost straight; insertion of fin base well forward

of pelvic-fin origin; spine origin over pectoral-fin inner marg�n; sp�ne base broad, exposed anter�orly well below junction of spine and fin; shorter than second dorsal-fin spine; pre-first dorsal fin length 3.43 (3.17–3.70) in TL; first dorsal-fin length 2.16 (1.78–2.36) times its height, 1.26 (1.10–1.38) times second dorsal-fin length; first dorsal-fin height 1.51 (1.42–1.71) times second dorsal-fin height; first dorsal spine broken in holotype, often damaged but usually about half of fin height in other material. Second dorsal fin smaller than first, strongly to weakly raked; subtr�angular �n holotype and a few large spec�mens (�.e. NMNZ P 39908) w�th broadly rounded apex, weakly concave poster�or marg�n and long free rear tip; most other specimens retain fin form similar to juven�les, be�ng relat�vely taller, more strongly raked w�th a deeply concave poster�or marg�n, and shorter free rear tip; inner margin length 0.89 (0.84–1.20) times fin height; second dorsal-fin length 2.57 (2.16–2.88) times �ts he�ght; second dorsal sp�ne broken �n holotype, sp�ne length 0.7–1.2 in fin height in other material; fin-spine origin well behind free rear tip of pelvic fin, exposed below level of junction with spine and fin; second spine w�th broad base, slender d�stally, taper�ng above po�nt of

Figure 3. Cusps of the flank denticles of: A. Squalus griffini holotype (NMNZ P 662, female 972 mm TL); and B. Squalus griffini (NMNZ P 20965, female 482 mm TL). F�eld of v�ew w�dth 0.8 mm.

A

B


Table 1. Proport�onal d�mens�ons as percentages of total length for the holotype (NMNZ P 662) and ranges for the 16 paratypes of Squalus griffini. Note that those measurements �n �tal�cs are hor�zontal rather than d�rect measurments.

S. griffini Holotype Paratypes

M�n. Max.TL – Total length 972 473 1070PCL – Precaudal length 80.2 76.8 80.3PD2 – Pre-second dorsal length 63.2 59.4 63.2PD1 – Pre-first dorsal length 29.1 27.0 31.5SVL – Pre-vent length 52.2 50.0 54.9PP2 – Prepelv�c length 49.5 47.0 52.0PP1 – Prepectoral length 20.1 19.3 24.4HDL – Head length 21.0 20.2 24.3PG1 – Prebranch�al length 18.2 16.5 20.7PSP – Presp�racular length 12.1 10.0 14.5POB – Preorb�tal length 7.5 7.4 8.8PRN – Prenar�al length 5.2 5.0 5.9POR – Preoral length 9.5 8.8 11.4INLF – Inner nostr�l-lab�al furrow space 4.4 4.1 5.5MOW – Mouth w�dth 5.8 5.7 7.6ULA – Lab�al furrow length 2.2 1.8 2.7INW – Internar�al space 4.5 4.4 5.2INO – Interorb�tal space 7.1 7.3 9.6EYL – Eye length 3.6 3.9 5.3EYH – Eye he�ght 1.4 1.4 2.4SPL – Sp�racle length 1.5 1.0 1.8GS1 – F�rst g�ll-sl�t he�ght 2.3 1.4 2.3GS5 – F�fth g�ll-sl�t he�ght 2.1 1.7 2.5IDS – Interdorsal space 25.3 22.6 26.0DCS – Dorsal-caudal space 9.7 9.1 11.1PPS – Pectoral-pelv�c space 26.9 21.5 27.0PCA – Pelv�c-caudal space 23.8 21.7 26.0D1L – F�rst dorsal length 14.9 13.3 16.5D1A – F�rst dorsal anter�or marg�n 11.5 11.5 14.6D1B – F�rst dorsal base length 9.7 8.3 10.4D1H – F�rst dorsal he�ght 6.9 6.3 8.7D1I – F�rst dorsal �nner marg�n 5.2 4.9 6.4D1P – F�rst dorsal poster�or marg�n 8.2 7.5 10.4D1ES – F�rst dorsal sp�ne length – 1.9 4.0D1BS – F�rst dorsal sp�ne base w�dth 0.8 0.6 0.9D2L – Second dorsal length 11.8 10.7 13.1D2A – Second dorsal anter�or marg�n 9.5 9.1 11.6D2B – Second dorsal base length 7.7 6.7 8.1D2H – Second dorsal he�ght 4.6 4.2 5.8D2I – Second dorsal �nner marg�n 4.1 3.8 5.1D2P – Second dorsal poster�or marg�n 5.4 4.3 6.3D2ES – Second dorsal sp�ne length – 3.1 5.3D2BS – Second dorsal sp�ne base w�dth 0.7 0.6 0.9

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Table 1. cont’d.

S. griffini Holotype Paratypes

M�n. Max.P1A – Pectoral anter�or marg�n 13.6 13.4 16.5P1I – Pectoral �nner marg�n 6.5 5.4 7.7P1B – Pectoral base length 5.3 5.0 6.2P1P – Pectoral poster�or marg�n 9.7 9.5 13.2P2L – Pelv�c length 10.2 9.0 11.5P2H – Pelv�c he�ght 4.7 4.4 5.1P2I – Pelv�c �nner marg�n 4.2 4.7 5.7CDM – Dorsal caudal marg�n 19.4 19.4 22.5CPV – Preventral caudal marg�n 10.0 9.8 11.7CPU – Upper postventral caudal marg�n 14.8 14.7 17.2CPL – Lower postventral caudal marg�n 3.2 3.4 4.8CFW – Caudal fork w�dth 6.2 6.0 7.1CFL – Caudal fork length 8.4 8.5 10.1HANW – Head w�dth at nostr�ls 6.5 6.4 7.5HAMW – Head w�dth at mouth 10.4 10.4 12.4HDW – Head w�dth 11.6 12.2 14.9TRW – Trunk w�dth 11.7 8.3 13.7ABW – Abdomen w�dth 7.9 8.0 14.0TAW – Ta�l w�dth 6.0 5.3 7.2CPW – Caudal peduncle w�dth 3.1 2.1 4.8HDH – Head he�ght 8.8 7.9 11.0TRH – Trunk he�ght 9.4 8.6 13.9ABH – Abdomen he�ght 9.0 6.9 14.7TAH – Ta�l he�ght 6.7 4.9 6.8CPH – Caudal peduncle he�ght 2.3 2.2 2.6CLO – Clasper outer length – 0.8 4.6CLI – Clasper �nner length – 5.0 9.7CLB – Clasper base w�dth – 0.8 1.4

exposure, generally reaching level of insertion of fin when unbroken (sl�ghtly poster�or �n some); �nterdorsal space 0.79 (0.76–1.07) �n pre-pectoral length, 1.15 (1.12–1.30) in pre-first dorsal fin length. Pectoral fin large, anterior marg�n weakly convex, apex rounded; poster�or marg�n concave to nearly stra�ght; �nner marg�n convex, length 6.5 (5.4–7.7)% TL, free-rear t�p narrowly rounded, not acute; base 2.59 (2.35–2.85) �n anter�or marg�n length. Pelvic fins small, anterior and posterior margins almost stra�ght, apex broadly rounded, free rear t�p narrow, acute. Precaudal tail tapering to caudal fin, broadly sem�c�rcular poster�orly, ventral groove well developed; prom�nent lateral keels extend�ng poster�orly from below insertion of second dorsal fin past caudal fin insertion; pelv�c–caudal space 1.13 (0.84–1.22) �n pectoral–pelv�c space, 0.85 (0.76–1.07) �n pre-pectoral length; dorsal–

caudal space 2.61 (2.10–2.68) �n �nterdorsal space; upper and lower precaudal pits present, upper pit better defined than lower pit. Caudal fin well developed, dorsal caudal marg�n 19.4 (19.4–22.5)% TL, 1.08 (0.94–1.19) �n head length, w�thout a subterm�nal notch; lower caudal lobe 1.93 (1.72–2.03) �n dorsal caudal marg�n. Vertebral centra 121 (113–119), monospondylous 47 (45–47), precaudal 91 (86–90) and caudal 30 (26–29). Teeth m�ss�ng from upper and lower jaws of holotype, 11 + 8 rema�n�ng �n upper jaw, 8 + 10.5 rema�n�ng �n the lower jaw; 13–14+13=26–27 �n upper jaw, 10–12+10–12=21–24 �n lower jaw of other mater�al (NMNZ P 2759, NMNZ P 22549, NMNZ P 35270, NMNZ P 35973 and NMNZ P 41774).

COLOUR.— The female holotype �s sta�ned brown and


Figure 4. Lateral view of the dorsal fins of Squalus griffini: A. first dorsal fin, and C. second dorsal fin of NMNZ P 39893 (female 1018 mm TL); B. first dorsal fin, and D. second dorsal fin of CSIRO H 6006–11 (�mmature male 473 mm TL).

has lost all natural colour and mark�ngs. Large female w�th s�m�lar morphology to holotype (NMNZ P 39893, 1018 mm TL, based on fresh spec�men): un�form grey-brown dorsally, wh�te below; dorsal and ventral colours strongly demarcated on head, extend�ng from snout below eye and through gill slits to pectoral-fin base, demarcation less distinct posteriorly; dorsal and caudal fins grey, first dorsal-fin anterior base and free rear tip paler than rest of fin, second dorsal-fin anterior base not darker than rest of fin, dorsal-fin tips and upper posterior margin with narrow black margin; first dorsal spine pale, second dorsal spine dark brown to dusky at base, becom�ng pale towards t�p; pectoral, pelvic and caudal fins grey with white posterior margins and tips; naked axils of fins and pectoral origin dusky; eyes br�ght green �n l�fe. Some �nd�v�duals w�th

scattered black spots on dorsal and ventral surfaces. A melan�st�c spec�men (NMNZ P 5176) �s black above, dark grey below w�th wh�te spots scattered along the ventral surface from the mouth to the pelvic fins. Juvenile female (NMNZ P 41775, 418.5 mm TL): first and second dorsal-fin anterior bases and free rear tips distinctly paler than rest of body and fin, posterior margin of second dorsal-fin white almost to tip; first dorsal-fin upper anterior and upper posterior margins and tip black; second dorsal-fin upper anterior margin and tip black; caudal fin with broad wh�te poster�or marg�n, no caudal bar; basal half of upper caudal lobe w�th th�n black fr�nge, upper caudal blotch fa�nt, d�stal th�rd of lobe wh�te; d�stal half of lower caudal lobe wh�te. Some adult spec�mens reta�n�ng juven�le colour pattern. SIZE.— Females reach at least 1100 mm TL (spec�men not reta�ned) and males at least 898 mm TL (NMNZ P 39908); smallest mature male 693 mm TL (NMNZ P 41774); smallest pregnant female 865 mm TL, largest �mmature female 935 mm TL (spec�mens not reta�ned).

DISTRIBUTION.— North and South Islands of New Zealand and Chatham R�se north of the Subtrop�cal Front. Also occurs north of New Zealand on Wanganella Bank, Norfolk and Lou�sv�lle R�dges, and southern Kermadec R�dge to at least Raoul Island.

REMARKS.— Garr�ck (1960) synonym�sed S. griffini w�th S. blainville (syn. S. blainvillii, S. blainvillei) based on the l�terature and an exam�nat�on of the holotype of S. griffini, several spec�mens from New Zealand, and a presumed S. blainville spec�men from the east coast of the Un�ted States. Use of the name S. blainville �s problemat�c as there are no extant types and R�sso’s (1827) description and figure do not conform to any known spec�es of Squalus (Chen et al. 1979; Muñoz-Chápul� and Ramos 1989). Consequently two reg�onal taxonom�c rev�ews have descr�bed morpholog�cally d�sparate spec�es from Japan and the eastern North Atlant�c and Med�terranean as S. blainville. Although recogn�s�ng that R�sso’s descr�pt�on was problemat�c, Chen et al. (1979) nonetheless identified a Japanese species as S. blainville on the basis of its high dorsal fins and long fin spines also depicted in Risso’s figure. Muñoz-Chápuli and Ramos (1989) however observed that the only Squalus spec�es known from the Med�terranean Sea (type local�ty of S. blainville) w�th long sp�nes agrees well w�th the�r �nterpretat�on (erroneously) of S. megalops. Hence, rather than recogn�s�ng th�s spec�es as S. blainville, they identified it as S. megalops, and appl�ed S. blainville to a less abundant short-sp�ned spec�es �n order to ma�nta�n nomenclatural stab�l�ty w�th B�gelow and Schroeder (1948, 1957), Garr�ck (1960) and Bass et al. (1976), who had appl�ed th�s name to short-sp�ned spec�es from the Southern Hem�sphere. Muñoz-Chápul� and Ramos (1989) concluded that the Japanese spec�es referred by Chen et al. (1979) to S. blainville �s probably undescr�bed.

A

B

C

D

98

Notw�thstand�ng the problems regard�ng correct appl�cat�on of the name Squalus blainville identified by Muñoz-Chápul� and Ramos (1989), S. griffini �s d�st�ngu�shable from ‘S. blainville’ from Japan on the basis of its much smaller first dorsal fin (height 6.4–8.7%, ma�nly less than 7.9% vs. 9.1–10.3% TL), more delicate dorsal-fin spines, and the first dorsal-fin base greater than �ts he�ght (rather than less than �ts he�ght). Squalus griffini �s d�st�ngu�shable from S. blainville from the eastern North Atlant�c and Med�terranean by �ts longer snout (preoral length 8.8–11.4, mean 9.5% TL vs. 8.4–8.7% TL �n S. blainville) and smaller first dorsal fin (first dorsal-fin height mainly less than 7.9%, mean 6.9% TL vs. 8.1–8.5% TL). Squalus griffini �s morpholog�cally most s�m�lar to the short-sp�ned ‘m�tsukur�� group’ sensu Chen et al. (1979) w�th�n what was formerly known as the ‘fernand�nus-bla�nv�lle group’ (B�gelow and Schroeder 1948, 1957; Garr�ck 1960). Members of the ‘m�tsukur�� group’ are characterised by having tricuspid denticles, pectoral fins not falcate w�th the�r �nner marg�n rounded rather than pointed, a short first dorsal-fin spine, and relatively high vertebral counts (Chen et al. 1979; Muñoz-Chápul� and Ramos 1989). However, S. griffini �s read�ly d�st�ngu�shed from members of the ‘m�tsukur�� group’ by the absence of a dark caudal bar. It d�ffers from S. mitsukurii �n hav�ng a lower vertebral count (113–121, mean 116 vs. 118–127, mean 121) and generally larger �nterdorsal space (distance 1.12–1.30 times in pre-first dorsal fin length vs. 1.45–1.73) (Chen et al. 1979; Last et al. 2007b, Part 6 of th�s �ssue). The absence of a dark caudal bar also separates S. griffini from the two Austral�an spec�es, S. chloroculus Last, Wh�te and Motomura, 2007 and S. montalbani Wh�tley, 1931, prev�ously referred to S. mitsukurii. These spec�es also have a smaller second dorsal fin with a proportionally longer free rear tip, and much lower vertebral counts than S. griffini. Squalus

montalbani also has a shorter prenar�al snout (4.0–5.3% vs. 5.0–5.9%, mean 5.4% TL �n S. griffini) (Last et al. 2007b, Part 6 of th�s �ssue).

F�ve other Squalus spec�es w�th characters typ�cal of the ‘fernand�nus-bla�nv�lle group’ are descr�bed from Austral�a �n th�s �ssue. Squalus grahami Wh�te, Last and Stevens, 2007 (Part 7 of th�s �ssue) �s read�ly d�st�ngu�shed from S. griffini by the presence of a dark upr�ght caudal bar, fewer monospondylous vertebrae (38–42 vs. 45–47 �n S. griffini), a longer second dorsal-fin base and w�der prenar�al snout. Squalus edmundsi Wh�te, Last and Stevens, 2007 (Part 7 of th�s �ssue), S. albifrons Last, Wh�te and Stevens, 2007 (Part 5 of th�s �ssue) and S. altipinnis Last, Wh�te and Stevens, 2007 (Part 5 of th�s �ssue), all resemble S. griffini �n hav�ng a predom�nately wh�te poster�or caudal marg�n (S. edmundsi has an oblique caudal bar that barely reaches the posterior fin marg�n). However, unl�ke S. griffini, they have a more erect first dorsal fin with a longer, fatter spine (exposed base w�dth 0.9–1.2% vs. 0.6–0.9% TL, mean 0.7% TL �n S. griffini) resembl�ng that of S. blainville sensu Chen et al. (1979). Squalus edmundsi and S. altipinnis have fewer monospondylous vertebrae (42–44), and S. altipinnis has a shorter snout, smaller �nternar�al space and longer caudal peduncle than S. griffini. Squalus albifrons also d�ffers �n hav�ng a shorter snout, hor�zontal prenar�al length 2.91–3.06 t�mes �n preoral length (vs. 2.37–2.65 t�mes �n S. griffini), first dorsal-fin height 1.67–1.77 time �n �ts length (vs. 1.78–2.36 t�mes), and a sl�ghtly longer upper caudal-fin lobe (dorsal caudal margin 21.3–23.6% vs. 19.4–22.5%, mean 21% TL). Squalus notocaudatus Last, White and Stevens, 2007 also has an upright first dorsal fin with a relatively heavy spine (0.7–1.0% TL) but �s most read�ly d�st�ngu�shed from S. griffini by �ts colorat�on (d�agonal black caudal bar across base of lower caudal), short snout (preorb�tal length 6.4–7.0% vs. 7.4–8.8% TL), longer upper caudal (23.3–24.0% TL) and h�gh vertebral count (123–127 total vertebrae).

The holotype of Squalus griffini �s qu�te d�fferent �n body shape to most spec�mens collected from the reg�on. Th�s morphotype, represented �n the NMNZ collect�on by a few large �nd�v�duals, �s rather robust w�th broad, weakly tr�cusp�d dent�cles, a relat�vely small, subtr�angular second dorsal fin with a long free rear tip, and is possibly dark bod�ed. The more typ�cal form of S. griffini caught �n the reg�on �s less robust, has strongly tr�cusp�d dent�cles, and a relatively tall, deeply notched second dorsal fin. Colour of the body and fins usually becomes darker and more un�form w�th growth, however some adults rema�n relatively pale overall, with a pale first dorsal-fin anterior base, prominent dark fin markings, and whitish dorsal-fin free rear tips and lower caudal-fin lobe. Molecular work supports conspecificity of these morphotypes (see Part 12 of th�s �ssue, Ward et al., 2007) and no obv�ous morphometr�c or mer�st�c d�fferences ex�st between them. However, g�ven such rad�cal d�fferences �n the morphology of the denticles, and fin shape and colour,

Figure 5. Juvenile coloration of the caudal fin of Squalus griffini (CSIRO H 6066–08, �mmature male 262 mm TL).


th�s spec�es requ�res further �nvest�gat�on across �t d�str�but�onal range.

ACKNOWLEDGEMENTS

The project was part�ally funded by the New Zealand B�od�vers�ty Strategy, and the B�osystemat�cs of New Zealand EEZ F�shes program FRST Contract MNZ0302 to Te Papa. Cl�ve Roberts (Museum of New Zealand, Te Papa Tongarewa) supported a research v�s�t by the jun�or author to work on th�s problem. Kelly Tarlton’s Underwater World prov�ded collect�ng opportun�t�es �n the Bay of Plenty, and Bert Lee (Tolga Bay East Cape Charters) and Malcolm Franc�s, Mart�n Cryer and M�chael Mann�ng (NIWA) ass�sted w�th the collect�on of spec�mens from poorly sampled locat�ons. Andrew Stewart and Carl Struthers (Museum of New Zealand, Te Papa Tongarewa) prov�ded �nvaluable ass�stance w�th collect�on management, and the measurement and photography of specimens. Tess Fitzwater, Pacific Rad�ology, and Thomas Schultz, Te Papa, rad�ographed spec�mens. Dent�cle samples were prepared by Adr�an Turner, School of B�olog�cal Sc�ences, and SEMed w�th the help of John Montgomery at the School of Eng�neer�ng, Un�vers�ty of Auckland. John Pogonosk� and W�ll�am Wh�te (CSIRO) prov�ded techn�cal ass�stance and ed�tor�al comments.

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Part 10 — Squalus hemipinnis sp. nov, a new short-snout spurdog from eastern Indonesia

W�ll�am T. Wh�te, Peter R. Last and Gordon K. Yearsley


ABSTRACT.— A new spec�es of spurdog, Squalus hemipinnis sp. nov., �s descr�bed based on spec�mens from eastern Indones�a. Th�s spec�es d�ffers from other Squalus spec�es �n the reg�on �n a comb�nat�on of coloration, meristics, and morphometrics of the head, trunk and fins. Squalus hemipinnis does not conform to any of the w�dely recogn�sed Squalus subgroups, �nstead shar�ng characters w�th several groups. Squalus hemipinnis has a short snout typified by members of the ‘megalops-cubensis group’ but also has a dark caudal bar �n juven�les typ�cal of the ‘m�tsukur�� group’. Its low vertebral count �s unusual w�th�n Squalus.

Key words. Squalo�dea – Squal�dae – Squalus hemipinnis – new spec�es – Indones�a


INTRODUCTION

Market surveys at var�ous land�ng s�tes �n eastern Indones�a between Apr�l 2001 and March 2006 produced a w�de var�ety of new or poorly known sharks, skates, rays and ch�maeras, �nclud�ng undescr�bed squal�ds (Wh�te et al., 2006). Amongst th�s mater�al were four members of the genus Squalus, �nclud�ng one part�cularly abundant spec�es that �s very d�st�nct from �ts congeners and does not conform to any of the recogn�sed Squalus complexes. Th�s spec�es, wh�ch �s can be d�st�ngu�shed from sympatr�c relat�ves by the comb�nat�on of a relat�vely short snout, deeply forked second dorsal fin and the presence of a broad caudal bar �n juven�les, �s descr�bed based on ava�lable Indones�an mater�al. Compar�sons are made w�th other spec�es of Squalus from the Indo–West Pacific.

METHODS

Methods follow those outl�ned �n Part 1 of th�s �ssue (Last et al., 2007). Both morphometr�cs and mer�st�cs were taken from the holotype (MZB 15040) and the follow�ng 5 paratypes: CSIRO H 5631–02, CSIRO H 5631–06, CSIRO H 5692–02, CSIRO H 5692–03 and CSIRO H 5692–06 (Table 1). In add�t�on, mer�st�cs were taken from the follow�ng 9 paratypes: CSIRO H 5631–01, CSIRO H 5631–03, CSIRO H 5631–04, CSIRO H 5692–01, CSIRO H 5857–13, CSIRO H 5857–14, CSIRO H 5857–15, CSIRO H 5693–06 and CSIRO H 5693–07, and colour �s descr�bed from the follow�ng paratype embryo: CSIRO H 5889–38 (1 of 3). In the descr�pt�on, morphometr�c and mer�st�c values for the holotype are

given first followed in parentheses by the ranges of the paratypes.

Type spec�mens are depos�ted �n the �chthyolog�cal collect�on of the Museum Zoolog�cum Bogor�ense, Jakarta (MZB) and at the Austral�an Nat�onal F�sh Collect�on, Hobart (CSIRO); other mater�al of the new spec�es �s depos�ted �n the Nat�onal Museum of V�ctor�a (NMV); their registration numbers are prefixed with these acronyms.

Squalus hemipinnis sp. nov.


Squalus sp. 3: Wh�te et al., 2006, Economically Important Sharks and Rays of Indonesia, pp 70–71.

Holotype. MZB 15040, female 637 mm TL, Kedonganan fish landing site, Bali, Indonesia, 08°45′ S, 115°10′ E, July 2002.Paratypes. 17 spec�mens. Collected at same local�ty as holotype. CSIRO H 5631–01, female 580 mm TL; CSIRO H 5631–02, female 575 mm TL; CSIRO H 5631–03, female 605 mm TL; CSIRO H 5631–04, female 580 mm TL; CSIRO H 5631–06, female 566 mm TL; CSIRO H 5692–01, adult male 418 mm TL; CSIRO H 5692–02, adult male 498 mm TL; CSIRO H 5692–03, adult male 484 mm TL; CSIRO H 5692–06, female 566 mm TL; CSIRO H 5693–06, female 550 mm TL; CSIRO H 5693–07, female 425 mm TL; CSIRO H 5857–13, female 640 mm TL; CSIRO H 5857–14, female 660 mm TL; CSIRO H 5857–15, female 625 mm TL;

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CSIRO H 5889–38, 3 female embryos 162–166 mm TL.Non-types. 9 spec�mens. Collected at same local�ty as holotype. MZB 15039, adult male 493 mm TL; NMV A 29559–001, adult male 452 mm TL; NMV A 29559–002, female 580 mm TL; CSIRO H 5631–05, female 564 mm TL; CSIRO H 5692–05, four embryos 154–157 mm TL; CSIRO H 5889–29, adult male 449 mm TL.

DIAGNOSIS.— A moderately s�zed spec�es of Squalus w�th the follow�ng comb�nat�on of characters: body slender, depth 9.0–10.6% TL; snout relat�vely narrow, very short, hor�zontal prenar�al length 2.0–2.2 t�mes mouth w�dth, preoral length 2.41–2.49 t�mes hor�zontal prenar�al length, 8.4–9.1% TL; eye large, �ts length 5.1–5.6% TL; secondary lobe of anterior nasal flap well developed; dorsal fins small, strongly raked; first dorsal-fin spine moderate, broad-based; second dorsal-fin spine moderate, robust and broad-based; pectoral fin of adult weakly falcate; pectoral-fin inner margin relatively short, 6.9–7.8% TL; caudal fin with a short, broad caudal bar, enlarged upper caudal blotch and fr�nge located d�stally on lobe in juveniles; flank denticles broadly unicuspidate to weakly tr�cusp�date; 35–38 monospondylous centra, 72–76 precaudal centra, 96–100 total centra; adult max�mum s�ze at least 78 cm TL.

DESCRIPTION.— Body fus�form, slender, nape prom�nently humped (less so �n smallest paratypes); deepest near first dorsal-fin spine, maximum depth 1.14 (0.86–1.09 �n 5 paratypes) t�mes w�dth; trunk depth 1.05 (0.84–0.98) t�mes abdomen depth; head short 22.6 (20.9–22.5)% TL; caudal peduncle slender, 27.1 (26.4–28.4)% TL. Head not espec�ally broad, w�dth 1.30 (1.03–1.18) t�mes trunk w�dth, 1.27 (1.15–1.30) t�mes abdomen w�dth; depressed forward of sp�racles, becom�ng somewhat subtriangular towards pectoral-fin origin; length 2.09 (2.15–2.37) �n pre-vent length; he�ght 0.85 (0.78–0.89) t�mes w�dth. Snout short, narrowly tr�angular �n lateral v�ew, apex bluntly po�nted, broadly po�nted �n dorsal v�ew; lateral prenar�al marg�n not angular; hor�zontal length 1.07 (1.05–1.14) t�mes eye length, 0.72 (0.70–0.74) t�mes �nterorb�tal space; hor�zontal prenar�al length 2.49 (2.41–2.47) �n preoral length. Eye oval, large, length 4.23 (4.04–4.19) �n head, 2.34 (2.10–2.72) t�mes �ts he�ght; strongly notched poster�orly, notch deep anter�orly, usually weakly connected to anteroventral marg�n of sp�racle. Sp�racle s�ze moderate, broadly crescent�c (somet�mes subtr�angular); broad lobe-l�ke fold on poster�or marg�n; greatest d�ameter 3.04 (2.98–3.33) �n eye length. G�ll open�ngs d�rected sl�ghtly anterodorsally from bottom to top (more upr�ght in some paratypes); first four subequal in size, fifth longest, height of fifth slit 2.0 (1.9–2.3)% TL. Mouth almost transverse, upper jaw weakly concave, w�dth 1.15 (1.14–1.22) �n preoral length; upper lab�al furrows about 1.5 t�mes length of lower furrows; prom�nent postoral groove, subequal �n length to upper lab�al furrows, extend�ng posterolaterally from angle of jaws; two ser�es of funct�onal teeth �n upper jaw, three (somet�mes two)

ser�es �n lower jaw. Teeth s�m�lar �n upper and lower jaws; upper teeth un�cusp�d, �nterlock�ng, blade-l�ke, cusps d�rected strongly laterally, low; tooth base broader than length of �ts cusp. Nostr�ls small, almost transverse; anterior nasal flap strongly bifurcate; anterior lobe broad, posterior lobe much shorter, narrow, somewhat flattened; �nternar�al space 2.29 (2.10–2.42) �n preoral length, 2.69 (2.38–2.56) t�mes nostr�l length. Dermal dent�cles (based on holotype) on flank very small, non-imbricate; crowns well elevated, quadrate, broadly un�cusp�date to weakly tr�cusp�date w�th pronounced med�an r�dge; med�an r�dge commenc�ng very well anter�or of rest of crown, w�th a mes�al furrow develop�ng anter�orly and converg�ng rap�dly towards poster�or t�p of crown; poster�or port�on of cusp strongly produced, pungent; lateral port�on of crown very short, cusps weakly developed or form�ng an obtuse angle; c�rcumorb�tal reg�on eas�ly abraded �n holotype (and most paratypes). First dorsal fin small, strongly raked, narrowly rounded ap�cally (somet�mes angular); anter�or marg�n moderately convex; upper poster�or marg�n almost stra�ght, not vert�cal, �nstead d�rected very sl�ghtly anterodorsally from bottom to top (vert�cal �n most paratypes), weakly concave near free rear t�p (var�able, never strongly concave); free rear tip very thick basally, short; inner margin of fin almost stra�ght; �nsert�on of base extremely well forward of pelvic-fin origin, well posterior to free rear tip of pectoral fin; fin-spine origin slightly posterior to pectoral-fin �nsert�on; sp�ne base broad, exposed anter�orly well below junction of spine and soft portion of fin; soft portion of fin connected well above mid-point of total sp�ne length (somet�mes at about three-quarters of sp�ne length from �ts base); sp�ne taper�ng sl�ghtly d�stally, anter�or marg�n almost stra�ght; exposed port�on raked; spine longer than exposed portion of second dorsal-fin sp�ne (var�able �n paratypes, d�st�nctly shorter �n CSIRO H 5692–03), exposed base more elevated than exposed base of second dorsal-fin spine; pre-first dorsal length 3.52 (3.42–3.61) times in TL; first dorsal-fin length 1.89 (1.73–2.15) t�mes �ts he�ght, 1.15 (1.09–1.25) t�mes second dorsal-fin length; first dorsal-fin height 2.10 (1.77–1.91) times second dorsal-fin height; exposed first dorsal spine length 0.60 (0.55–0.73) times height of fin. Second dorsal fin very small, very strongly raked; anter�or marg�n moderately convex (less so �n some paratypes), apex narrowly angular; poster�or marg�n very deeply concave, max�mum concav�ty form�ng a V-shape (mostly less than 90º) at about m�d-po�nt of marg�n, upper port�on d�rected dorsoposter�orly very strongly from bottom to top; free rear t�p moderately elongate, �nner margin length 1.09 (1.03–1.18) times fin height; second dorsal-fin length 3.45 (2.94–3.22) times its height; spine length 1.01 (1.26–1.41) in height of fin; fin-spine origin well behind free rear tip of pelvic fin, exposed at about or sl�ghtly below level of junct�on w�th sp�ne and soft portion of fin; exposed second spine broad based (broader than exposed base of first dorsal-fin spine); spine robust, bluntly po�nted d�stally, taper�ng rap�dly just above po�nt of exposure, sp�ne t�p not extend�ng to level of �nsert�on


Figure 1. Lateral v�ew of: A. Squalus hemipinnis sp. nov. holotype (MZB 15040, female 637 mm TL); B. Squalus hemipinnis (CSIRO H 5631–02, female 575 mm TL).

Figure 2. Ventral v�ew of the head of Squalus hemipinnis sp. nov. holotype (MZB 15040, female 637 mm TL).

of fin (extending beyond insertion in paratypes); �nterdorsal space 0.97 (0.85–0.91) �n length from snout tip to pectoral-fin origin, 1.19 (1.11–1.16) in pre-first dorsal length; �nterdorsal r�dge weak (appear�ng as a shallow groove in some paratypes). Pectoral fin moderate (relat�vely smaller �n smallest paratypes), anter�or marg�n moderately convex; �nner marg�n moderately convex, length 7.8 (6.9–7.3)% TL; apex narrowly rounded, lobe-

l�ke, weakly falcate (not falcate �n smallest paratypes); poster�or marg�n moderately concave (otherw�se weakly), free rear tip broadly angular; fin base very short, 3.18 (2.82–3.11) in length of anterior margin. Pelvic fins size moderate, anter�or and poster�or marg�ns almost stra�ght, apex broadly rounded, free rear t�p broadly angular (more acute �n mature males). Caudal peduncle very long, tapering slightly to caudal fin; subcircular in cross-section

A

B

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Table 1. Proport�onal d�mens�ons as percentages of total length for the holotype (MZB 15040) and ranges for the 5 paratypes of Squalus hemipinnis sp. nov.

S. hemipinnis sp. nov. Holotype Paratypes

M�n. Max.TL – Total length 637 484 575PCL – Precaudal length 79.4 79.3 80.0PD2 – Pre-second dorsal length 60.4 60.3 62.6PD1 – Pre-first dorsal length 28.4 27.7 29.2SVL – Pre-vent length 47.3 47.3 49.6PP2 – Prepelv�c length 45.8 45.5 48.3PP1 – Prepectoral length 22.7 21.2 22.0HDL – Head length 22.6 20.9 22.5PG1 – Prebranch�al length 18.8 17.1 18.9PSP – Presp�racular length 11.7 10.6 12.4POB – Preorb�tal length 6.6 6.4 6.7PRN – Prenar�al length 4.3 3.9 4.4POR – Preoral length 8.8 8.4 9.1INLF – Inner nostr�l-lab�al furrow space 5.2 4.8 5.3MOW – Mouth w�dth 7.6 6.9 7.7ULA – Lab�al furrow length 2.0 1.9 2.1INW – Internar�al space 3.8 3.5 4.3INO – Interorb�tal space 7.9 7.6 8.3EYL – Eye length 5.3 5.1 5.6EYH – Eye he�ght 2.3 1.9 2.6SPL – Sp�racle length 1.8 1.5 1.8GS1 – F�rst g�ll-sl�t he�ght 1.9 1.7 2.0GS5 – F�fth g�ll-sl�t he�ght 2.0 1.9 2.3IDS – Interdorsal space 23.9 24.2 25.9DCS – Dorsal-caudal space 10.6 10.9 11.4PPS – Pectoral-pelv�c space 20.9 20.9 24.4PCA – Pelv�c-caudal space 27.1 26.4 28.4D1L – F�rst dorsal length 14.4 12.8 15.3D1A – F�rst dorsal anter�or marg�n 13.0 11.7 13.8D1B – F�rst dorsal base length 9.3 8.1 9.9D1H – F�rst dorsal he�ght 7.7 6.7 7.4D1I – F�rst dorsal �nner marg�n 5.4 4.9 5.4D1P – F�rst dorsal poster�or marg�n 8.2 6.8 8.2D1ES – F�rst dorsal sp�ne length 4.6 4.1 5.5D1BS – F�rst dorsal sp�ne base w�dth 0.9 0.7 0.8D2L – Second dorsal length 12.6 11.4 12.5D2A – Second dorsal anter�or marg�n 11.4 10.7 11.6D2B – Second dorsal base length 8.8 7.4 8.2D2H – Second dorsal he�ght 3.6 3.8 3.9D2I – Second dorsal �nner marg�n 4.0 4.1 4.6D2P – Second dorsal poster�or marg�n 3.7 3.6 4.4D2ES – Second dorsal sp�ne length 3.7 4.8 5.5D2BS – Second dorsal sp�ne base w�dth 1.0 0.9 1.0


Table 1. cont’d.

S. hemipinnis sp. nov.

Holotype ParatypesM�n. Max.

P1A – Pectoral anter�or marg�n 15.7 13.4 14.8P1I – Pectoral �nner marg�n 7.7 6.9 7.3P1B – Pectoral base length 4.9 4.4 5.1P1P – Pectoral poster�or marg�n 11.6 9.5 10.1P2L – Pelv�c length 10.0 9.0 10.1P2H – Pelv�c he�ght 5.2 4.2 5.0P2I – Pelv�c �nner marg�n 3.3 3.9 4.8CDM – Dorsal caudal marg�n 20.6 19.5 20.7CPV – Preventral caudal marg�n 11.3 11.1 11.8CPU – Upper postventral caudal marg�n 15.3 13.7 15.1CPL – Lower postventral caudal marg�n 5.8 4.6 6.0CFW – Caudal fork w�dth 7.1 6.1 6.9CFL – Caudal fork length 8.7 8.6 9.3HANW – Head w�dth at nostr�ls 6.9 6.5 7.6HAMW – Head w�dth at mouth 9.3 8.9 10.0HDW – Head w�dth 12.1 10.5 12.0TRW – Trunk w�dth 9.3 9.4 10.6ABW – Abdomen w�dth 9.5 8.5 9.6TAW – Ta�l w�dth 6.6 5.7 6.5CPW – Caudal peduncle w�dth 2.9 2.5 2.9HDH – Head he�ght 10.2 8.5 9.7TRH – Trunk he�ght 10.6 9.0 10.2ANH – Abdomen he�ght 10.2 9.1 11.8TAH – Ta�l he�ght 7.0 6.1 7.0CPH – Caudal peduncle he�ght 2.2 2.2 2.5CLO – Clasper outer length – 4.5 4.5CLI – Clasper �nner length – 7.1 7.3CLB – Clasper base w�dth – 1.4 1.5

anter�orly, broadly sem�c�rcular poster�orly; ventral groove well developed; lateral keels well developed, or�g�nat�ng sl�ghtly poster�or to (or below) �nsert�on of second dorsal fin, terminating about half an eye diameter behind caudal-fin insertion; pelvic–caudal space 0.77 (0.73–0.92) �n pectoral–pelv�c space, 0.84 (0.77–0.81) �n prepectoral length; dorsal–caudal space 2.27 (2.12–2.38) �n �nterdorsal length; dorsal caudal p�t well developed, ventral caudal pit rudimentary. Caudal fin short, upper postventral marg�n almost stra�ght (weakly convex �n some paratypes), apex of lower lobe narrowly angular; dorsal caudal marg�n 1.10 (1.03–1.13) �n head length; length of lower caudal lobe 1.83 (1.67–1.80) �n upper lobe length. Vertebral centra 96 (96–100 �n 14 paratypes), monospondylous 36 (35–38), precaudal 72 (72–76) and caudal 24 (22–26). Teeth �n upper jaw (�n paratype

CSIRO H 5692–06) 13+13=26, lower jaw 12+11=23.

COLOUR.— When fresh (based on paratype CSIRO H 5631–02): slate grey above, wh�te below; l�ght and dark tonal areas sharply demarcated on head, extend�ng from snout adjacent suborb�t, across head and through top part of g�ll sl�ts; d�fferent�at�on �nd�st�nct on trunk. F�rst dorsal fin greyish, darker apical marking appearing as a weak blotch rather than a marg�nal mark�ng; anter�or base and extrem�ty of rear t�p sl�ghtly paler; sk�n-covered basal portion of spine and base of soft portion of first dorsal fin white; second dorsal-fin anterior base white, rest of fin grey�sh w�th a small prom�nent black blotch d�stally, free rear tip pale; dorsal-fin spines dusky with darker anterior margins. Caudal fin mainly greyish with a broad white poster�or marg�n; pale edge extend�ng along full length

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of poster�or marg�n, from near apex of upper lobe to near apex of lower lobe, �ts w�dth constr�cted sl�ghtly at caudal fork; sl�ghtly darker black caudal str�pe present; no obv�ous caudal bar or other prom�nent black blotches or mark�ngs. In preservat�ve (based on holotype): s�m�lar to fresh paratype except fins distinctly more brownish than body; �nterface between l�ght and dark tonal areas extends through mid-gill area; pectoral and pelvic fins brownish grey above w�th broad wh�te poster�or marg�ns; pectoral and pelvic fins brownish below with broad pale posterior marg�ns and wh�te bases; upper caudal str�pe ev�dent, no other dark mark�ngs. Late-term embryo (CSIRO H 5889–38, female 166 mm TL) w�th strongly d�fferent�ated l�ght and dark fin markings; light and dark tonal areas across head sharply defined. First dorsal fin with a prominent white anterior base and free rear tip, distal part of fin mostly black; second dorsal fin with oblique black bar extending from base to fin apex, anterior portion of fin base and free rear t�p wh�te; prom�nent med�an black stripe extending from insertion of second dorsal fin to origin of caudal fin. Caudal fin with a prominent black caudal bar at �ts fork; bar short extend�ng from angle of fin diagonally over posterior half of base of ventral lobe, intersection with posterior fin margin short; upper caudal fr�nge pronounced, closer to t�p of lobe than �ts or�g�n; upper caudal blotch well defined, located in central distal portion of upper lobe near fin margin; most of lower lobe and posterior margin of upper lobe white. Pectoral fin with dark anter�or marg�n and basal blotch; broad poster�or white margin; fin more uniformly pale ventrally. Pelvic fins uniformly white dorsally and ventrally. SIZE.— Females and males reach at least 780 and 522 mm TL, respect�vely; smallest mature male 432 mm TL, smallest post-natal male 395 mm TL; smallest post-natal female 325 mm TL; 6 pregnant females exam�ned had between 7 and 10 late-term embryos, 140–175 mm TL.

Figure 3. Cusps of the flank denticles of Squalus hemipinnis sp. nov. paratype (CSIRO H 5631–06, female 566 mm TL). F�eld of v�ew w�dth 1.0 mm.

Figure 4. Lateral view of the dorsal fins of: Squalus hemipinnis sp. nov. paratype (CSIRO H 5631–02, female 575 mm TL) – A. first dorsal fin, B. second dorsal fin.

DISTRIBUTION.— Known only from eastern Indones�a between C�lacap �n Central Java (ca. 08°S, 109°E) and Tanjung Luar �n eastern Lombok (ca. 09°S, 117°E) where �t �s the most abundant squalo�d targeted by demersal longline fishers. Depth distribution unknown as existing material was collected from fish markets in eastern Indones�a, but l�kely to be deeper than 100 m.

ETYMOLOGY.— Der�ved from the comb�nat�on of the Greek hemi (half) and the Lat�n pinna (fin) in allusion to �ts strongly notched, v-shaped poster�or marg�n of the second dorsal fin.

VERNACULAR.— Indones�an Shortsnout Spurdog.

REMARKS.— Squalus hemipinnis can be d�st�ngu�shed from all other members of the genus by the comb�nat�on of a short ‘megalops-l�ke’ snout, deeply notched second dorsal-fin posterior margin, strongly demarcated dorsal colorat�on, very low number of precaudal centra, and other morphometr�c and colour deta�ls. Squalus hemipinnis has a fewer precaudal centra (72–75) than other Squalus spec�es (more than 78), except S. acanthias L�nnaeus, 1758 (74–79 �n Austral�an spec�mens) and S. lalannei Baranes, 2003 (67–69). Squalus hemipinnis d�ffers from S. acanthias �n hav�ng a shorter snout, more anteriorly positioned dorsal fin, no white spots on the dorsal and lateral body surfaces, and a b�furcate nasal flap (vs. a single-lobed nasal flap). Although S. lalannei

A

B


was not �ncluded �n the recent book Sharks of the World (Compagno et al., 2005), �t d�ffers from all other Squalus spec�es �n hav�ng an unusually low number of precaudal centra, �.e. 67–69 (vs. 72–75 �n S. hemipinnis).

The snout shape of S. hemipinnis �s s�m�lar to members of the ‘megalops-cubensis group’ and ‘highfin megalops group’ but �t �s clearly separable from all members of these Squalus subgroups. It possesses low, strongly raked dorsal fins (first dorsal-fin height 6.7–7.7% TL), which differ from the taller, more upright dorsal fins of S. albifrons Last, Wh�te and Stevens, 2007 (7.7–8.9% TL), S. altipinnis Last, Wh�te and Stevens, 2007 (7.8–7.9% TL), S. bucephalus Last, Séret and Pogonosk�, 2007 (8.1–8.5% TL), S. crassispinus Last, Edmunds and Yearsley, 2007 (7.2–8.9% TL) and S. notocaudatus Last, Wh�te and Stevens, 2007 (8.2–9.4% TL). Squalus hemipinnis d�ffers morphometr�cally from S. cubensis Howell R�vero, 1936, �n hav�ng a shorter snout (preorb�tal length 6.4–6.7 vs. 7.2–7.6% TL; preoral length 1.14–1.21 vs. about 1.44 t�mes mouth w�dth, 8.4–9.1 vs. 9.6–10.8% TL), shorter pectoral-fin free rear tips (pectoral-fin inner marg�n 1.90–2.09 vs. 1.42–1.55 �n �ts anter�or marg�n) and a shorter second dorsal-fin free rear tip (second dorsal-fin inner margin 1.03–1.18 vs. 1.42–1.44 times �ts he�ght). Squalus hemipinnis d�ffers from S. megalops (Macleay, 1881) �n hav�ng a narrower head (head w�dth at anter�or mouth 8.9–10.0 vs. 10.9–12.8% TL, head w�dth 10.5–12.1 vs. 12.8–14.7% TL), a short, constr�cted snout (hor�zontal prenar�al length 2.02–2.17 vs. 2.58–3.40 times mouth width) and more robust dorsal-fin spines. The caudal fin of S. hemipinnis, wh�ch has a d�st�nct black bar on �ts fork �n juven�les that becomes unobv�ous in adults, differs from the northwestern Pacific species, S. brevirostris Tanaka, 1917, which has a pale caudal-fin poster�or marg�n w�thout a dark caudal bar.

Although S. hemipinnis �s morpholog�cally s�m�lar to some members of the ‘megalops-cubens�s group’, �t possesses

Figure 5. Juvenile coloration of the caudal fin of Squalus hemipinnis sp. nov. paratype (CSIRO H 5889–38, female embryo 166 mm TL).

a d�st�nct caudal bar �n juven�les typ�cal of members of the ‘m�tsukur�� group’ and the ‘japon�cus group’ but lack�ng �n the ‘megalops-cubens�s group’. Squalus hemipinnis d�ffers from members of the ‘m�tsukur�� group’ and the ‘japon�cus group’ �n hav�ng a shorter snout (preoral length 8.4–9.1% TL vs. >11.4 % TL �n S. japonicus Ish�kawa, 1908 and S. nasutus Last, Marshall and Wh�te, 2007, and 9.3–11.5% �n S. chloroculus Last, Wh�te and Motomura, 2007, S. edmundsi Wh�te, Last and Stevens, 2007, S. grahami Wh�te, Last and Stevens, 2007, S. mitsukurii Jordan and Snyder �n Jordan and Fowler, 1903 and S. montalbani Wh�tley, 1931) and a strongly notched, v-shaped second dorsal-fin posterior margin. Squalus griffini Ph�ll�pps, 1931, has a much longer snout (preoral length 7.4–8.8% TL vs. 6.4–6.7 % TL), more strongly tr�cusp�d dent�cles, and a d�fferent caudal colorat�on.

ACKNOWLEDGEMENTS

Market surveys of eastern Indones�a were conducted as part of a 5-year ACIAR funded project. Spec�al thanks go Junaedi (Kedonganan fish market) who retained many spec�mens for us, �nclud�ng the holotype, between our regular v�s�ts to Indones�a, and Fahm� (LIPI), Dharmad� (RCCF) and Jenny G�les (QDPI&F) for the�r ass�stance in the field. Thanks also go to Alastair Graham for assembl�ng collect�on mater�al, Mart�n Gomon and D� Bray (NMV) for supply�ng reg�strat�on deta�ls, Lou�se Conboy for ed�t�ng �mages of the types and other mater�al, Bryn Farmer for supply�ng dent�cle �mages, T�m Founta�n for prepar�ng rad�ographs and obta�n�ng mer�st�c �nformat�on, Dan Gledh�ll for ass�st�ng w�th a l�terature summary, John Stevens for obta�n�ng tooth counts and prov�d�ng ed�tor�al comments, John Pogonosk� for techn�cal ass�stance and ed�tor�al comments. Other �mportant contr�butors �n th�s project �nclude Subhat Nurhak�m (RCCF), Ono Kurnaen Sumadh�harga (LIPI), Ian Potter (Murdoch Un�vers�ty), and Steve Blaber and Cathy D�chmont (CSIRO).

REFERENCES

Baranes, A. (2003) Sharks from the Am�rantes Islands, Seychelles, w�th a descr�pt�on of two new spec�es of squalo�ds from the deep sea. Israel Journal of Zoology, 49, 33–65.


Howell R�vero, L. (1936) Some new, rare and l�ttle-known fishes from Cuba. Proceedings of the Boston Society of Natural History, 41, 41–76.

Ish�kawa, C. (1908) Descr�pt�on of a new spec�es of

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squalo�d shark from Japan. Proceedings of the Academy of Natural Sciences of Philadelphia, 60, 71–73.

Jordan, D.S. and Fowler, H.W. (1903) A rev�ew of the elasmobranchiate fishes of Japan. Proceedings of the United States National Museum, 26, 593–674.

Last, P.R., Edmunds, M. and Yearsley, G.K. (2007) Part 2 — Squalus crassispinus sp. nov., a new spurdog of the ‘megalops-cubens�s group’ from the eastern Ind�an Ocean, p. 11–22. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Last, P.R., Marshall, L.J. and Wh�te, W.T. (2007) Part 8 — Squalus nasutus sp. nov, a new long-snout spurdog of the S. japonicus group from the Ind�an Ocean, p. 83–90. In: Descriptions of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Last, P.R., Séret, B. and Pogonosk�, J.J. (2007) Part 3 — Squalus bucephalus sp. nov., a new short-snout spurdog from New Caledon�a, p. 23–29. In: Descr�pt�ons of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.

Last, P.R., Wh�te, W.T. and Motomura, H. (2007) Part 6 — A descr�pt�on of Squalus chloroculus sp. nov., a new spurdog from southern Austral�a, and the resurrect�on of S. montalbani Wh�tley, p. 55–69. In: Descr�pt�ons of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.








Wh�te, W.T., Last, P.R. and Stevens, J.D. (2007) Part 7 — Two new spec�es of Squalus of the ‘m�tsukur�� group’ from the Indo–Pacific, p. 71–81. In: Descr�pt�ons of new dogfishes of the genus Squalus (Squalo�dea: Squal�dae). Eds: Last, P.R., Wh�te, W.T. and Pogonosk�, J.J., CSIRO Marine and Atmospheric Research Paper 014, 130 pp.



Part 11 — Clarification of the status of Squalus tasmaniensis and a diagnosis of Squalus acanthias from Australia, including a key

to the Indo–Australasian species of Squalus

W�ll�am T. Wh�te, Gordon K. Yearsley and Peter R. Last


ABSTRACT.— Squalus tasmaniensis has been prev�ously synonym�sed w�th S. megalops and S. mitsukurii. However, recent exam�nat�on of the holotype revealed that �t �s a juven�le S. acanthias based on the pos�t�on of the first dorsal fin and the narrow head. Squalus acanthias can be separated from all other known spec�es of Squalus by the posterior location of the first dorsal fin (its origin behind the free rear tips of pectoral fins), a single-lobed anterior nasal flap, and the presence of white spots on the dorsal and lateral body surfaces. A d�agnos�s of southeastern Austral�an populat�ons of S. acanthias and a key to the Indo–Australas�an Squalus spec�es are also presented.

Key words. Squalo�dea – Squal�dae – Squalus tasmaniensis – Squalus acanthias – key to spec�es – Indo–Australas�an.


INTRODUCTION

Squalus tasmaniensis Howell R�vero, 1936a was descr�bed based on a s�ngle juven�le spec�men collected off Hobart, Tasman�a, �n 1860. The author noted �n h�s descr�pt�on that the spec�men “resembles Squalus acanthias L�nné in many aspects, and was identified as such by Garman”. Its nomenclatural affinity has remained unclear, having been synonym�sed w�th S. mitsukurii Jordan and Snyder, 1903 (Compagno, 1984) and w�th S. megalops (Macleay, 1881) by Last and Stevens (1994). Recent exam�nat�on of the poorly preserved holotype of S. tasmaniensis by one of the authors (P. Last) revealed that th�s juven�le type spec�men resembles S. acanthias L�nnaeus, 1758 �n a number of key characters.

Myagkov and Kondur�n (1986) proposed that S. acanthias compr�sed at least 4 subspec�es, S. a. acanthias from the northern Atlant�c, S. a. africana from southern Afr�ca, S. a. ponticus from the Black Sea and S. a. subsp. from Austral�a and New Zealand. However, the descr�pt�ons prov�ded by Myagkov and Kondur�n (1986) are very short, often based on embryos, and rel�ed heav�ly on characters that are subjected to allometr�c changes w�th growth. Compagno et al. (1991) synonym�sed the subspec�es S. a. africana w�th S. acanthias, but S. a. ponticus �s cons�dered a val�d subspec�es by Eschmeyer (2006). Apparently restr�cted to the Black Sea, S. a. ponticus, �s much larger than other forms of

S. acanthias, atta�n�ng lengths of up to 180 cm TL (vs. <100 cm) w�th the females and males matur�ng at greater than 104 (vs. <60) and 120 (vs. <70) cm TL, respect�vely (Myagkov and Kondur�n, 1986; Compagno et al., 2005).

Th�s paper proposes the synonymy of S. tasmaniensis w�th S. acanthias and prov�des a d�agnos�s of S. acanthias from southeastern Austral�a. A key to the Indo–Austral�an spec�es of Squalus �s also �ncluded.

METHODS

Methods follow those outl�ned �n Part 1 of th�s publ�cat�on (Last et al., 2007). The holotype of S. tasmaniensis (MCZ 146-S) and five southeastern Australian specimens of S. acanthias (CSIRO T 712, CSIRO H 1214, CSIRO H 4226–01, CSIRO H 4876–01 and CSIRO H 6485–01) were measured �n full (Table 1). For compar�son w�th the juven�le (240 mm TL) holotype of S. tasmaniensis, selected measurements were also taken from s�m�lar-s�zed, juven�le/late term embryos of S. acanthias (n=4), S. chloroculus (n=3) and S. megalops (n=3) �n the 200-271 mm TL s�ze range. Mer�st�cs were taken from rad�ographs of 9 spec�mens of S. acanthias. Spec�mens exam�ned are depos�ted �n the Austral�an Nat�onal F�sh Collect�on, Hobart (CSIRO) and the �chthyolog�cal collect�on of the Museum of Comparat�ve Zoology, Boston (MCZ); the�r registration numbers are prefixed with these acronyms.

110

Squalus acanthias L�nnaeus, 1758

F�gs 1, 2, Table 1

Squalus acanthias L�nnaeus, 1758, Systema Naturae, ed. 10, 1: 233. Poss�ble syntypes (2): L�nnaean Collect�on �n Uppsala, no. 159 (alcohol), no. 160 (dr�ed), European Seas (“Hab�tat �n Oceano Europaeo”).

Material examined. 15 spec�mens. MCZ 146-S (Squalus tasmaniensis holotype), female 240 mm TL, Hobart, Tasman�a; CSIRO H 1214, female 678 mm TL, Dover or Southport, Tasman�a, 30 m; CSIRO H 2921–01, female 606 mm TL, CSIRO H 2921–02, adolescent male 525 mm TL, CSIRO H 2921–03, female 698 mm TL, off Shark Point, Tasmania, 42°48′ S, 147°29′ E, 5.5 m; CSIRO H 4226–01, female 429 mm TL, Freder�ck Henry Bay, Tasmania, 42°50′ S, 147°33′ E, 6 m; CSIRO H 6205–01, �mmature male 271 mm TL, Battery Po�nt, Tasmania, 42°53′ S, 147°20′ E; CSIRO T 1099, 4 late-term embryos 205–218 mm TL, Sandy Bay, Tasman�a, 42°53′ S, 147°20′ E; CSIRO H 4876–01, adult male 616 mm TL, off Woodbridge, Tasmania, 43°10′ S, 147°15′ E, 20 m; CSIRO T 712, adult male 575 mm TL,

CSIRO H 6485–01, female 492 mm TL, CSIRO T 783, adult male 661 mm TL, off Port Davey, Tasman�a.

DIAGNOSIS.— A moderately s�zed spec�es of Squalus w�th the follow�ng comb�nat�on of characters: body very slender, abdomen w�dth 7.2–9.2% TL; head narrow, w�dth at mouth 7.9–10.2% TL; mouth w�dth 1.7–2.2 t�mes hor�zontal prenar�al length; snout moderately long, preoral length 2.2–2.5 t�mes hor�zontal prenar�al length, 8.5–9.8% TL; anterior nasal flap single-lobed; dorsal fins small and raked, first dorsal-fin height 1.1–1.3 times its inner margin length, second dorsal-fin height 0.7–0.9 times its inner margin length; first dorsal-fin origin located just posterior to pectoral-fin free rear tips; exposed bases of dorsal-fin spines relatively narrow, 0.4–0.7% TL; first dorsal-fin spine short, exposed length 1.7–2.7% TL; pectoral-fin anterior margin 1.9–3.1 times �ts �nner marg�n length; preventral caudal marg�n 1.9–2.6 times inner margin of pelvic fin; caudal fin pale with poorly demarcated, wh�t�sh marg�n, black�sh caudal blotch at apex of upper lobe, anter�or marg�ns of both lobes wh�t�sh �n juven�les; no dark caudal bar; dorsal and lateral surfaces of body blu�sh grey w�th an �rregular array of moderately-large white spots; flank denticles tricuspid;

Figure 1. Lateral v�ew of: A. Squalus acanthias (CSIRO H 1214, female 678 mm TL); B. Squalus tasmaniensis (MCZ 146–S, female 247 mm TL); C. Squalus acanthias (CSIRO H 6205–01, �mmature male 271 mm TL).

A

B

C


Table 1. Proport�onal d�mens�ons as percentages of total length for 5 southeastern Austral�an spec�mens of Squalus acanthias and the holotype (MCZ 146-S) of S. tasmaniensis.

S. tasmaniensis S. acanthias (n = 5) Holotype M�n. Max.

TL – Total length 240 429 678PCL – Precaudal length 78.2 77.7 79.1PD2 – Pre-second dorsal length 56.3 58.8 61.0PD1 – Pre-first dorsal length 29.5 32.8 35.2SVL – Pre-vent length 49.3 50.6 52.4PP2 – Prepelv�c length 46.2 48.7 50.8PP1 – Prepectoral length 19.7 20.7 22.9HDL – Head length 20.1 20.7 22.4PG1 – Prebranch�al length 15.7 17.6 19.2PSP – Presp�racular length 10.6 10.8 12.0POB – Preorb�tal length 6.1 6.8 7.2PRN – Prenar�al length 3.2 4.2 5.0POR – Preoral length 7.5 8.5 9.8INLF – Inner nostr�l-lab�al furrow space 3.9 3.9 4.6MOW – Mouth w�dth 6.2 7.0 8.7ULA – Lab�al furrow length – 1.5 2.6INW – Internar�al space – 3.3 4.0INO – Interorb�tal space – 6.8 8.5EYL – Eye length 3.5 3.4 3.9EYH – Eye he�ght 1.0 1.4 2.2SPL – Sp�racle length 1.3 0.9 1.4GS1 – F�rst g�ll-sl�t he�ght 1.7 1.4 1.9GS5 – F�fth g�ll-sl�t he�ght 1.1 1.7 2.4IDS – Interdorsal space 18.7 19.8 21.2DCS – Dorsal-caudal space 12.3 10.8 12.0PPS – Pectoral-pelv�c space 21.2 21.3 24.9PCA – Pelv�c-caudal space 22.2 21.5 23.0D1L – F�rst dorsal length 13.5 12.4 13.1D1A – F�rst dorsal anter�or marg�n 12.0 9.3 10.2D1B – F�rst dorsal base length 8.1 7.0 7.8D1H – F�rst dorsal he�ght 4.3 5.9 6.8D1I – F�rst dorsal �nner marg�n 5.6 5.0 5.7D1P – F�rst dorsal poster�or marg�n 5.6 6.5 8.4D1ES – F�rst dorsal sp�ne length 1.9 1.7 2.7D1BS – F�rst dorsal sp�ne base w�dth 0.5 0.4 0.6D2L – Second dorsal length 14.0 12.3 13.3D2A – Second dorsal anter�or marg�n 10.4 9.2 10.6D2B – Second dorsal base length 9.3 7.5 8.4D2H – Second dorsal he�ght 3.0 3.2 4.1D2I – Second dorsal �nner marg�n 4.9 4.8 5.0D2P – Second dorsal poster�or marg�n 5.2 4.7 5.7D2ES – Second dorsal sp�ne length 3.7 2.6 3.7D2BS – Second dorsal sp�ne base w�dth 1.0 0.6 0.7

112

Table 1. cont’d.

S. tasmaniensis S. acanthias (n = 5) Holotype M�n. Max.

P1A – Pectoral anter�or marg�n 13.7 13.6 16.2P1I – Pectoral �nner marg�n 7.0 4.4 7.3P1B – Pectoral base length 4.8 4.7 6.0P1P – Pectoral poster�or marg�n 8.5 8.1 11.0P2L – Pelv�c length 10.7 10.0 11.3P2H – Pelv�c he�ght 3.2 4.2 5.0P2I – Pelv�c �nner marg�n 3.8 4.1 5.6CDM – Dorsal caudal marg�n 22.1 20.0 21.9CPV – Preventral caudal marg�n 12.4 10.6 10.9CPU – Upper postventral caudal marg�n 14.7 14.3 15.1CPL – Lower postventral caudal marg�n 4.7 3.2 4.6CFW – Caudal fork w�dth 6.2 6.6 7.6CFL – Caudal fork length 10.8 9.1 10.7HANW – Head w�dth at nostr�ls 6.6 5.6 6.7HAMW – Head w�dth at mouth – 7.9 10.2HDW – Head w�dth 10.1 10.2 12.3TRW – Trunk w�dth 6.4 8.2 11.4ABW – Abdomen w�dth 4.2 7.2 9.2TAW – Ta�l w�dth – 5.2 5.8CPW – Caudal peduncle w�dth – 2.2 2.9HDH – Head he�ght 5.5 8.4 9.6TRH – Trunk he�ght – 8.8 10.3ABH – Abdomen he�ght – 8.6 10.9TAH – Ta�l he�ght 3.5 5.2 6.3CPH – Caudal peduncle he�ght 2.1 2.3 2.7CLO – Clasper outer length – 5.6 6.2CLI – Clasper �nner length – 10.7 11.1CLB – Clasper base w�dth – 1.2 1.3

41–45 monospondylous centra, 74–79 precaudal centra, 100–105 total centra; adult max�mum s�ze at least 100 cm TL.

REMARKS.— The holotype of Squalus tasmaniensis (F�g. 1b) resembles members of the ‘acanth�as group’ �n having a more posteriorly located first dorsal fin compared to all other Squalus species, i.e. the first dorsal-fin origin �s s�tuated just poster�or to (rather than over or forward of) the pectoral-fin free rear tips and the interdorsal space �s short (�nterdorsal space 18.7% TL �n holotype, 17.9–18.6% TL �n s�m�lar-s�zed S. acanthias vs. 20.7–23.9% TL �n s�m�lar-s�zed S. chloroculus Last, Wh�te & Motomura, 2007 and S. megalops). The head of the holotype of S. tasmaniensis �s also much narrower (F�g. 1b) than that of e�ther S. chloroculus or S. megalops, w�th wh�ch �t has

prev�ously been synonym�sed (head w�dth 10.1% TL �n holotype, 10.9–11.5% TL �n s�m�lar-s�zed S. acanthias vs. 12.5–14.2% TL �n s�m�lar-s�zed S. chloroculus and S. megalops). Thus, although the holotype of S. tasmaniensis �s �n a poor cond�t�on, �t shares a number of key features w�th S. acanthias and g�ven �ts s�ze appears to be a juven�le of th�s spec�es. Th�s spec�men was originally identified by Garman as S. acanthias and the confus�on that has followed seems to be related to the lack of any obv�ous wh�te spots on the sk�n. The poor cond�t�on of the spec�men has also poss�bly contr�buted to its previous misidentification.

The genet�c component of th�s �ssue (Ward et al., 2007; Part 12) found that the Squalus acanthias samples separated �nto two clearly d�st�nct subgroups: a subgroup


consisting of 10 specimens from the North Pacific (Japan and U.S.A.); and larger subgroup from a much broader geograph�c range encompass�ng the Atlant�c (U.K., U.S.A. and Iceland) and South Pacific Oceans (Australia, New Zealand and Ch�le). Populat�ons belong�ng to the second subgroup are likely to conform to the current definition of S. acanthias because the holotype of S. acanthias was collected from European Seas. Add�t�onal morpholog�cal �nvest�gat�ons are requ�red to compare the reg�onal forms of S. acanthias, in particular North Pacific populations vs. Atlantic and South Pacific populations. There are several jun�or synonyms of S. acanthias from the Atlant�c (Acanthias americanus Storer, 1846, A. antiquorum Leach, 1818, Squalus barbouri Howell R�vero, 1936b, A. commun Navarrete, 1898, A. linnei Malm, 1877, Spinax mediterraneus G�stel, 1848, Spinax (Acanthias) suckleyi G�rard, 1855, A. vulgaris R�sso, 1827) and South Pacific (S. fernandinus Mol�na, 1782, S. kirki Ph�ll�pps, 1931, A. lebruni Va�llant, 1888, S. whitleyi Ph�ll�pps, 1931). Squalus wakiyae Tanaka, 1918 from Japan, wh�ch is the only available name from the North Pacific, may be resurrected as a val�d spec�es.

Other material. Squalus chloroculus: CSIRO H 1350–02, 4 embryos 217–238 mm TL, northwest of Macquar�e

Figure 2. Ventral v�ew of the head of A. Squalus acanthias (CSIRO H 4876–01, adult male 616 mm TL); B. Squalus tasmaniensis (MCZ 146–S, female 247 mm TL).

A

B

Harbour, Tasmania, 41°52′ S, 144°23′ E, 1370 m. Squalus megalops: CA 3176, �mmature female 255 mm TL, east of Gabo Island, New South Wales, 37°38′ S, 150° 14′ E, 580 m; CSIRO H 5320–01, immature male 200 mm TL, south-southeast of Cape Everard, Victoria, 38°09′ S, 149°38’ E, 294 m; CSIRO H 6449–01, �mmature male 245 mm TL, off Tr�al Harbour, Tasman�a, ca. 42° S, 145° E.

Key to Squalus of the Indo–Australasian region.

1a. First dorsal-fin spine origin distinctly behind free rear tips of pectoral fins; anterior nasal flap single-lobed; body typ�cally blu�sh grey w�th wh�te spots ............................S. acanthias (c�rcumglobal)

b. First dorsal-fin spine origin over or anterior to free rear tips of pectoral fins; anterior nasal flap with two lobes; body typ�cally grey�sh �n colour, lack�ng wh�te spots ............................................................ 2

2a. Snout long to very long, d�stance from snout t�p to �nner nostr�l longer than d�stance from �nner edge of nostr�l to lab�al furrow…..................3

b. Snout short, d�stance from snout t�p to �nner nostr�l shorter or equal to d�stance from �nner edge of nostr�l to lab�al furrow….....................................10

3a. Ventral lobe of caudal fin conspicuously black…..................................... S. melanurus (New Caledon�a)

b. Ventral lobe of caudal fin without a conspicuous black mark�ng ...................................................... 4

4a. First dorsal fin and its associated spine almost upr�ght ................................................................... 5

b. First dorsal fin and its associated spine angled rearwards ............................................................... 6

5a. Snout acutely po�nted, prenar�al length more than 6.5% TL; 46 monospondylous vertebrae ...............................S. rancureli (Vanuatu)

b. Snout narrowly tr�angular, prenar�al length less than 6.5% TL; 43–44 monospondylous vertebrae .. ............................................................S. edmundsi

(northwestern Austral�a, Indones�a) 6a. Snout narrow and long (preoral length 84–

103% of head w�dth); 36–42 (usually 37–40) monospondylous vertebrae ................................... 7

b. Snout relat�vely broad and long (preoral length 62–81% of head w�dth, rat�o larger �n spec�mens of S. griffini <550 mm TL); 41–47 (usually 43–47) monospondylous vertebrae ................................... 8

7a. Snout very long; preoral length <2.0 t�mes hor�zontal

prenarial length; undamaged first dorsal-fin spine much shorter than second dorsal-fin spine (first dorsal sp�ne length 74–90% of second dorsal sp�ne length); 78–84 (typ�cally 81 or fewer) precaudal

114

vertebrae ................................................. S. nasutus (northwestern Austral�a, Indones�a, Ph�l�pp�nes) b. Snout long; preoral length >2.3 t�mes hor�zontal

prenarial length; undamaged first dorsal-fin spine slightly shorter than second dorsal-fin spine (first dorsal sp�ne length about 96% of second dorsal sp�ne length); 80–87 (typ�cally 82 or more) precaudal vertebrae ................................................. ............................... S. grahami (eastern Austral�a)

8a. Caudal fin with a continuous broad pale posterior marg�n, lack�ng a dark caudal bar (most ev�dent �n juven�les and subadults; often �nd�st�nct �n largest adults); 86–91 (usually 87–90) precaudal vertebrae ......................... S. griffini (New Zealand)

b. Caudal fin with a dark caudal bar extending onto �ts poster�or marg�n (most ev�dent �n juven�les and subadults); 79–86 (usually 79–84) precaudal vertebrae ............................................................... 9

9a. Second dorsal fin moderately large, its length 12.2–13.9% TL, 4.2–4.9 �n pre-second dorsal length; adult claspers relat�vely long (outer length 4.5–5.6% TL) .......................................... S. montalbani

(Austral�a, Indones�a, Ph�l�pp�nes) b. Second dorsal fin relatively smaller, its length

10.9–12.2% TL, 5.0–5.8 �n pre-second dorsal length; adult claspers shorter (outer length 3.8–4.6% TL) ......... S. chloroculus (southern Austral�a)

10a. Dorsal body colorat�on strongly demarcated from ventral coloration; second dorsal-fin posterior marg�n strongly notched form�ng a d�st�nct v-shape; 100 or less total vertebrae (range 96–100) ............................... S. hemipinnis (Indones�a)

b. Dorsal body colorat�on only weakly demarcated from ventral coloration; second dorsal-fin posterior marg�n concave but not strongly notched and not form�ng a v-shape; 102 or more total vertebrae (range 102–127) ................................................. 11

11a. Head enlarged, very broad (head w�dth at anter�or of nostr�ls 7.7–9.3% TL); dent�cles on flank both unicuspid and weakly tricuspid in adults .................... S. bucephalus (New Caledon�a)

b. Head moderate, narrower (head w�dth at anter�or of nostr�ls 6.3–7.6% TL, usually less than 7.0% TL); denticles on flank either unicuspid or distinctly tr�cusp�d ............................................................. 12

12a. Prom�nent dark bar along base of lower caudal-fin lobe (most evident in juveniles); 123–127 total vertebrae .................................................................. ............... S. notocaudatus (northeastern Austral�a)

b. No dark oblique bar on base of lower caudal-fin lobe; usually less than 120 total vertebrae (range 102–122) ............................................................ 13

13a. First dorsal fin low (first dorsal-fin height 2.4–4.0%

TL), usually angled rearwards ........................... 14 b. First dorsal fin tall (first dorsal-fin height 4.4–5.9%

TL), usually upr�ght (somet�mes angled rearwards �n S. crassispinus) .............................................. 15

14a. Interdorsal space 24–25% TL; preoral length 1.9–2.2 t�mes �nternar�al space; 37–40 monospondylous vertebrae ............................ S. megalops (Austral�a)

b. Interdorsal space 23–24% TL; preoral length 2.3–2.5 t�mes �nternar�al space; 41–43 monospondylous vertebrae ................... S. raoulensis (New Zealand)

15a. Dorsal sp�nes very robust (second dorsal-sp�ne base width 1.3–1.5% TL); denticles on flank unicuspid; 107–111 total vertebrae ........................................... ................ S. crassispinus (northwestern Austral�a)

b. Dorsal sp�nes robust (second dorsal-sp�ne base width 0.7–0.9% TL); denticles on flank tricuspid; 114–122 total vertebrae ...................................... 16

16a. Caudal fin large (dorsal caudal margin >21% TL, <2.9 t�mes �n pre-second dorsal length); 44–46 (usually 45 or 46) monospondylous vertebrae ........ S. albifrons (northeastern Austral�a)

b. Caudal fin moderately large (dorsal caudal margin <20% TL, >3.0 t�mes �n pre-second dorsal length); 42–44 monospondylous vertebrae .......................... ..................... S. altipinnis (northwestern Austral�a)

ACKNOWLEDGEMENTS

S�ncere thanks go to Alasta�r Graham for assembl�ng collect�on mater�al, John Pogonosk� for ed�t�ng and techn�cal ass�stance, Lou�se Conboy for �mage work, and T�m Founta�n for tak�ng rad�ographs and obta�n�ng mer�st�c �nformat�on. Thanks also to Karsten Hartel (MCZ) for prov�d�ng access to type mater�al from Tasman�a.

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Part 12 — DNA barcoding discriminates spurdogs of the genus Squalus

Robert D. Ward1, Bronwyn H. Holmes1, Tyler S. Zemlak2 and Peter J. Sm�th3

1Wealth from Oceans Flagsh�p, CSIRO Mar�ne & Atmospher�c Research, GPO Box 1538, Hobart, TAS 7001, AUSTRALIA2Department of Integrat�ve B�ology, Un�vers�ty of Guelph, Guelph, Ontar�o, N1G 2W1, CANADA. Present address:

Department of B�ology, L�fe Sc�ence Centre, Dalhous�e Un�vers�ty, Hal�fax, Nova Scot�a, B3H 4J1, CANADA3NIWA, Pr�vate Bag 14901, Well�ngton, NEW ZEALAND

ABSTRACT.— S�xteen spec�es of Squalus were analysed (barcoded) for a 654 bp reg�on of the m�tochondr�al cytochrome c ox�dase I gene. These compr�sed four prev�ously recogn�zed spec�es (S. acanthias, S. brevirostris, S. japonicus and S. megalops), two resurrected spec�es (S. griffini and S. montalbani), e�ght new spec�es (S. albifrons, S. chloroculus, S. crassispinus, S. edmundsi, S. grahami, S. hemipinnis, S. nasutus and S. raoulensis) and two spec�es yet to be descr�bed, �nformally named here as the Squalus sp. (Ta�wan highfin) and the Squalus sp. (Lombok highfin). A total of 127 individuals were barcoded. Average sequence d�vergences w�th�n and between the 15 spec�es represented by mult�ple spec�mens were 0.17±0.05% and 4.35±0.23% respect�vely. Sequences of a part�cular spec�es always clustered t�ghtly together and away from clusters of other spec�es. No �nstances of spec�es shar�ng of sequences were observed, suggest�ng that these spec�es were reproduct�vely �solated from one another. DNA barcod�ng can be used to �dent�fy spec�mens of th�s genus w�th a h�gh degree of certa�nty.

Key words. Squal�dae – DNA barcode – m�tochondr�al DNA – cytochrome ox�dase – spec�es identification


Here we �nvest�gate whether DNA barcode analys�s can contr�bute to a better understand�ng of the taxonomy and identification of Squalus spec�es.

The taxonomy of Austral�an spurdogs was poorly known unt�l Last and Stevens (1994) recogn�sed 9 spec�es of Squalus �n Austral�an waters, 6 of wh�ch were prev�ously undescr�bed (spec�es A through F). The �dent�ty of the taxa found around northern New Zealand has also been uncerta�n and spec�mens have been var�ously (and �ncorrectly) recorded as Squalus mitsukurii (Paul�n et al., 2001; Tracey and Shearer, 2002) and Squalus blainville (Ayl�ng and Cox, 1982; Paul, 2000).

The preced�ng papers �n th�s spec�al publ�cat�on of the CSIRO Mar�ne & Atmospher�c Research Paper ser�es formally descr�be 11 spec�es from the Indo–Australas�an region. In this final paper of the series, we use the approach of DNA barcoding to confirm that these newly descr�bed Squalus spec�es do �ndeed have d�st�ngu�shable COI sequences cons�stent w�th the�r be�ng reproduct�vely �solated spec�es, and to support the concept that barcod�ng can be used for specimen identification in groups that are morphologically difficult to ascribe to species by non-experts. Some of these Squalus barcode data were briefly described in Ward et al. (2005), and subsequently �n Hauser (�n press), but �n those publ�cat�ons many of

INTRODUCTION

Molecular approaches have long been used to assist species identification, beginning with protein electrophores�s �n the early 1960s (Manwell and Baker, 1963) and m�tochondr�al DNA analyses �n the late 1970s (Av�se, 1994). A few years ago, �t was proposed that a s�ngle gene sequence m�ght be able to d�st�ngu�sh all, or the vast major�ty of, an�mal spec�es, and that the m�tochondr�al gene cytochrome c ox�dase I (COI) would be a good cand�date gene (Hebert et al., 2003). The spec�es sequence of COI was l�kened to a barcode (Hebert et al., 2003), although �t was recogn�sed from the start that spec�es m�ght each possess mult�ple sequences or barcodes. Th�s does not matter from a spec�es identification viewpoint providing that each sequence is un�que to an �nd�v�dual spec�es and clusters w�th (�s more closely related to) other sequences of that spec�es rather than w�th sequences from other spec�es.

DNA barcod�ng of more than 200 spec�es of Austral�an fishes (some imported) showed that COI barcoding does �ndeed perm�t the unamb�guous resolut�on of th�s assortment of fish species (Ward et al., 2005). Wh�le there may well be very closely related spec�es pa�rs that cannot be d�scr�m�nated by COI sequenc�ng, these are l�kely to account for no more than 1 or 2% of all spec�es.

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the spec�es rema�ned un-named. We also analyse barcode data from prev�ously descr�bed spec�es, �nclud�ng the w�despread S. acanthias.

METHODS

Deta�ls of the �nd�v�duals sequenced are prov�ded �n Table 1. S�xteen spec�es of Squalus were analysed, compr�s�ng 4 prev�ously recogn�sed spec�es (S. acanthias, S. brevirostris, S. japonicus and S. megalops), two resurrected spec�es (S. griffini and S. montalbani), 8 new spec�es (S. albifrons, S. chloroculus, S. crassispinus, S. edmundsi, S. grahami, S. hemipinnis, S. nasutus and S. raoulensis) and two spec�es yet to be described, informally identified here as the Squalus sp. (Taiwan highfin) and the Squalus sp. (Lombok highfin).

DNA was sequenced from wh�te muscle t�ssue samples stored at –80°C. DNA from most spec�mens was extracted us�ng a Chelex dry release procedure (s�m�lar to Haj�babae� et al., 2005), but New Zealand spec�mens were extracted us�ng a phenol-chloroform-ethanol procedure (Taggart et al., 1992). Approx�mately 655 base pairs (bp) were amplified from the 5’ region of the COI gene from m�tochondr�al DNA us�ng pr�mers F�shF2 and F�shR2 and PCR procedures as �n Ward et al. (2005), or pr�mers LCO1490 and HCO2198 descr�bed by Folmer et al. (1994) and amplifications using an initial denaturation of 95°C for 60 s; 30 cycles of 95°C for 30 s, 45°C for 60 s, and 72°C for 60 s, followed by a final extension at 72°C for 10 m�n. Products were labeled us�ng the B�gDye® Term�nator v3.1 Cycle Sequenc�ng K�t (Appl�ed B�osystems, Inc.) and sequenced b�d�rect�onally us�ng ABI 3730 or ABI 3100 cap�llary sequencers follow�ng manufacturer’s �nstruct�ons. Sequences were al�gned us�ng SeqScape v2.5 software (Appl�ed B�osystems, Inc.).

GenBank numbers, where depos�ted, are g�ven �n Table 1. We sequenced all �nd�v�duals except for a s�ngle S. acanthias sequence taken from GenBank. Th�s was a spec�men of S. acanthias taken off Iceland and whose ent�re mtDNA genome was sequenced (Rasmussen and Arnason, 1999, access�on number Y18134); the COI port�on of th�s sequence was added to our study.

Once sequences had been assembled for each spec�men, sequence d�vergences were calculated us�ng the K�mura two parameter (K2P) d�stance model (K�mura, 1980). Both ne�ghbour-jo�n�ng (NJ) and UPGMA trees of K2P d�stances were created to prov�de graph�c representat�ons of the pattern�ng of d�vergence between spec�es (Sa�tou and Ne�, 1987). Bootstrapp�ng was used to est�mate standard errors of sequence d�vergence and rel�ab�l�ty of tree branch po�nts, as performed �n MEGA3 (Kumar et al., 2004) w�th 1000 repl�cat�ons. The latter values are g�ven as percentages, and represent the percentage of t�mes that �nter�or branch �s computed �n the bootstrap resampl�ng techn�que.

RESULTS

A total of 127 spec�mens from 14 spec�es of Squalus (�nclud�ng those descr�bed �n the prev�ous papers of th�s �ssue) and two as-yet undescr�bed spec�es was barcoded us�ng the COI gene segment. The mean length of COI sequence across spec�mens was 645.5 bp, w�th a range of 573–654 bp. The average percentages of the four bases were thym�ne 33.8, cytos�ne 24.5, adenos�ne 25.1 and guan�ne 16.6.

The topology of the ne�ghbour-jo�n�ng K2P tree for all 127 sequences �s g�ven �n F�gs 1–4. All spec�mens of a part�cular spec�es clustered together and away from clusters of other spec�es. Note that one spec�men of S. grahami (BW–A094) had been erroneously identified as a S. mitsukurii �n Ward et al. (2005); subsequent morpholog�cal re-exam�nat�on revealed the �n�t�al error in identification.

F�gure 1 reveals three major Squalus clusters. Cluster A �s a large complex of twelve spec�es, w�th a bootstrap value of 95% (F�g. 2). It compr�ses one prev�ously-recogn�sed spec�es (S. japonicus), two resurrected spec�es (S. griffini and S. montalbani), 7 new spec�es (S. albifrons, S. chloroculus, S. crassispinus, S. edmundsi, S. grahami, S. hemipinnis and S. nasutus), and two presumpt�ve new spec�es that requ�re further study and descr�pt�on (�nformally known as Squalus sp. (Taiwan highfin) and Squalus sp. (Lombok highfin)). Cluster B comprises two prev�ously recogn�sed spec�es (S. brevirostris and S. megalops) and a new spec�es (S. raoulensis), w�th a bootstrap value of 100% (F�g. 3). Cluster C compr�ses a s�ngle prev�ously recogn�sed spec�es (S. acanthias), aga�n w�th a bootstrap value of 100% (F�g. 4).

Cluster A compr�ses 12 spec�es. F�fteen spec�mens of S. griffini (F�g. 2) from the north and south �slands of New Zealand formed a subcluster w�th a bootstrap value of 85% and a mean d�vergence of 0.03±0.02%. S�x spec�mens of S. grahami, all taken off the New South Wales coast of Austral�a, formed a subcluster w�th a bootstrap value of 60% and zero d�vergence. These two spec�es appeared to be qu�te closely related s�ster spec�es, form�ng a cluster w�th a bootstrap value of 95%. F�ve �nd�v�duals of S. japonicus were sequenced, all from Ta�wan. Two haplotypes were observed, wh�ch d�ffered �n a s�ngle synonymous mutat�on and appeared to make the taxon paraphylet�c. Average sequence d�vergence was 0.11±0.10%. Th�s spec�es grouped w�th S. nasutus form�ng a jo�nt cluster w�th 94% bootstrap support. E�ght spec�mens of S. nasutus (6 from Indones�a and two from an unrecorded locat�on, most l�kely Austral�a), formed a subcluster w�th 84% bootstrap support and a mean d�vergence of 0.18±0.13%. Two spec�mens of S. crassispinus, from northwestern Austral�a, had �dent�cal sequences and the pa�r had a bootstrap value


of 100%. N�ne spec�mens of S. montalbani, from the east and west coasts of Austral�a and from Indones�a, formed a subcluster w�th 76% bootstrap support and a mean d�vergence of 0.42±0.16%. Seven spec�mens of S. chloroculus, from the Great Austral�an B�ght to the Tasman Sea, formed a subcluster w�th 95% bootstrap support and a mean d�vergence of 0.11±0.07%. Fourteen spec�mens of S. edmundsi (F�g. 2), from Western Austral�a and Indones�a, formed a cluster w�th 65% bootstrap support and a mean d�vergence of 0.29±0.13%. Th�s spec�es cluster was pa�red (at 82% bootstrap support) w�th the S. hemipinnis spec�es cluster (F�g. 2). Th�s latter cluster compr�sed 6 spec�mens, all from Bal�, Indones�a, w�th 56% bootstrap support and a mean d�vergence of 0.19±0.10%. F�nally, there �s a cluster of three spec�es, Squalus albifrons and two undescr�bed spec�es, w�th a bootstrap support of 95%. F�ve spec�mens of S. albifrons were sequenced, all from New South Wales, Austral�a, wh�ch all had �dent�cal sequences form�ng a cluster w�th 81% bootstrap support. The two undescr�bed spec�es compr�sed two spec�mens of a Squalus sp. (Ta�wan highfin), with bootstrap support of 81% and a divergence of 0.19±0.18%, and a s�ngle spec�men of a Squalus sp. (Lombok highfin).

Cluster B compr�ses 3 spec�es. F�ve spec�mens of Squalus megalops, all taken from Bass Stra�t between V�ctor�a and Tasman�a (Austral�a), formed a subcluster w�th a bootstrap value of 60% and a mean d�vergence of 0.44±0.19% (F�g. 3). E�ght spec�mens of S. brevirostris, from southern Japan and Ta�wan, formed a subcluster w�th a bootstrap value of 89% and a mean d�vergence of 0.09±0.06%. Three spec�mens of S. raoulensis, from Raoul Island off the north of New Zealand, formed a subcluster w�th a bootstrap value of 93% and zero d�vergence.

Cluster C compr�ses the 31 �nd�v�duals of Squalus acanthias, a globally d�str�buted spec�es (F�g. 4). Ten of these were from the North Pacific (3 from Japanese waters, 7 from North Amer�ca) and 21 were from much of the rema�nder of �ts d�str�but�on (4 from the Un�ted K�ngdom – the Celt�c Sea, 5 from the Un�ted States – the Gulf of Ma�ne, one from Iceland, 5 from m�d-Ch�le – Isla Mocha, one from off the north �sland of New Zealand, and 5 from the east coast of Tasman�a). Spec�mens of S. acanthias had a mean sequence d�vergence of 0.47±0.16%. Two subclusters were evident, the first comprising the 10 north Pacific fish (bootstrap 60% and sequence d�vergence of 0.10±0.05%) and the other the 21 Atlantic/south Pacific fish (bootstrap 89% and sequence d�vergence of 0.14±0.05%). The �nter-subcluster divergence was 0.76±0.31%. One north Pacific fish (BW–A2328) d�ffered from all other S. acanthias �n hav�ng a tr�plet GAC rather than GAT about one-th�rd �nto the barcode reg�on. The GAC tr�plet �s found �n about 97.5% of all the other Squalus spec�mens barcoded, w�th GAT �n the res�dual 2.5%. Both tr�plets code for the same am�no ac�d, aspart�c ac�d.

Average sequence d�vergences w�th�n and between 15 spec�es were 0.17±0.05% and 4.35±0.23% respect�vely. Th�s calculat�on om�ts the Squalus sp. (Lombok highfin), for which thus far only one specimen has been barcoded.

Sequences were also clustered us�ng a UPGMA approach (F�gs 5–7), wh�ch assumes that the rate of nucleot�de subst�tut�on �s the same for all evolut�onary l�neages. Th�s assumpt�on �s not made by the NJ approach, except that the NJ tree shown here (F�g. 1) �s rooted at the m�dpo�nt of the longest route connect�ng two sequences. The UPGMA tree reta�ns all the major features descr�bed above from the NJ tree. One m�nor d�fference �s that �n the UPGMA tree, the S. japonicus spec�mens formed a s�ngle subcluster and no longer appeared paraphylet�c. Another d�fference �s that S. crassispinus now appeared basal w�th�n Cluster A.

DISCUSSION

All COI sequences, DNA barcodes, were found to be specific for particular species of Squalus. There was no ev�dence for shar�ng of barcodes among spec�es. However, the number of spec�mens barcoded per spec�es �s currently small, rang�ng from one to 31 (mean of 7.94), and undoubtedly new barcodes w�ll be descr�bed for these spec�es as sample s�zes �ncrease. Nevertheless, barcode var�at�on w�th�n spec�es �s currently very l�m�ted, and l�kely to rema�n so, and all barcodes thus far cluster by species. Hence, barcoded specimens can be identified to spec�es w�th a h�gh degree of certa�nty.

The lack of barcode shar�ng suggests that these spec�es of Squalus are reproduct�vely �solated from one another and th�s, together w�th the h�gh bootstrap values for the spec�es groups, supports the taxonom�c rev�s�ons descr�bed �n the earl�er papers of th�s �ssue.

Barcode analyses of some of these spec�es, pr�or to these taxonom�c rev�s�ons, were outl�ned �n F�gure 6 of Ward et al. (2005). In th�s earl�er paper, two clusters were labeled as S. mitsukurii – these are now known to be the resurrected spec�es S. montalbani and the new spec�es S. chloroculus. Further, one m�splaced S. mitsukurii �n Ward et al. (2005) was subsequently carefully reexam�ned morpholog�cally and found to be S. grahami – cons�stent with its barcode identification.

The two subclusters �n S. acanthias (north Pacific versus Atlantic and south Pacific) could suggest two cryptic species. In fact, the north Pacific form of S. acanthias was, pre-1960, given a different specific epithet, S. suckleyi (see Jensen, 1966). Hauser (�n press) po�nts out that wh�le the COI data show rec�procal monophyly cons�stent w�th distinct species status, some north Pacific specimens analysed �n h�s laboratory for the m�tochondr�al D-loop region fell into the north Atlantic/south Pacific cluster.

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Further work �s needed to resolve th�s �ssue, wh�ch should �nclude the analys�s of further spec�mens and perhaps nuclear DNA reg�ons.

The average COI sequence d�vergence between Squalus spec�es �n th�s study, 4.35%, was about 25 t�mes that w�th�n spec�es, 0.17%. These values are lower than the average d�stances between spec�es w�th�n genera (9.93%) and w�th�n spec�es (0.39%) recorded �n an exam�nat�on of 207 spec�es of elasmobranchs and teleosts (Ward et al., 2005), but the w�th�n-genus d�vergence �s very s�m�lar to a value of 3.6% found among 12 spec�es of the skate genus Bathyraja (Sp�es et al., 2006). Th�s low degree of var�at�on �s cons�stent w�th the reduced levels of molecular evolut�on detected w�th�n cart�lag�nous fishes (Martin and Palumbi, 1993). However, some teleosts show lower congener�c d�vergence, for example tunas of the genus Thunnus (1.11%, Ward et al., 2005). Thunnus spec�es may have rad�ated more recently than these Squalus spec�es.

The extremely h�gh, often complete, COI sequence �dent�ty between S. acanthias from the North Atlant�c (northeast and northwest), southeast Pacific (Chile), and southwest Pacific (Tasmania, New Zealand) suggests that these w�dely-d�vergent populat�ons are �n fact l�nked by present or recent gene flow, presumably through stepping-stone populat�on l�nks. In the western north Atlant�c, th�s spec�es undergoes seasonal m�grat�ons along the coast, from the Gulf of Ma�ne and Canad�an waters �n the summer to southern New England and North Carol�na �n the w�nter, and �n the east Atlant�c, from Br�ta�n’s northwest �n summer to Norway �n the w�nter (Jensen, 1966; McCauley et al., 2004). Wh�le S. acanthias �s more abundant �n temperate reg�ons, �ts d�str�but�on appears to extend throughout equator�al reg�ons so stepp�ng-stone gene flow is not unreasonable. Many marine fish – but not sharks – have pelag�c larval stages that can ass�st d�spersal. Squalus acanthias �s v�v�parous, w�th yolk-sac dependency, and has 1 to 32 young per l�tter (Compagno et al., 2005), and d�spersal �s l�kely to be med�ated largely by adult movements. Northern Pacific Ocean S. acanthias have been recorded as mak�ng extens�ve m�grat�ons, from Br�t�sh Columb�a to Japan (McFarlane and K�ng, 2003), so the genet�c homogene�ty observed between Japan and the North American Pacific coast is entirely expected.

We w�sh to emphas�se that barcod�ng �s essent�ally a tool for specimen identification rather than phylogenetic reconstruct�on, and can also reveal l�kely new spec�es. Its value in specimen identification became evident during the course of th�s work. The barcodes of several spec�mens were inconsistent with their original identifications, and subsequent morpholog�cal re-exam�nat�on of those voucher spec�mens revealed the�r true �dent�ty – �n every case cons�stent w�th the spec�es attr�buted by the barcode. Barcod�ng, at least w�th�n th�s genus, g�ves h�gher taxonom�c r�gour than that afforded by non-expert taxonom�sts. F�nally, w�th�n th�s paper we have descr�bed

related sequences as form�ng clusters, but we do not w�sh to �mply that the trees shown necessar�ly represent the true evolut�onary h�story of th�s group. Robust phylogenet�c reconstruct�ons need data from more than one gene reg�on, and preferably from both m�tochondr�al and nuclear genes. Hence the relat�onsh�ps and spec�es groups suggested here need to be confirmed (or refuted) w�th add�t�onal molecular data.

ACKNOWLEDGEMENTS

Th�s work would not have been poss�ble w�thout the expert taxonomists who provided specimen identification: Peter Last, Gordon Yearsley, Alasta�r Graham, W�ll�am Wh�te and Cl�nton Duffy. We thank Lorenz Hauser and J�m Franks from the Un�vers�ty of Wash�ngton for the t�ssue samples of Squalus acanthias from the north Pacific (Japan and USA), northwest and northeast Atlant�c (USA and UK) and southeast Pacific (Chile) and Hiroyuki Motomura from Kagosh�ma Un�vers�ty for samples of Squalus brevirostris from Japan. We thank Paul Hebert and the Un�vers�ty of Guelph team for sequenc�ng our first samples of Squalus and to John Pogonosk� and Gordon Yearsley for the�r ed�tor�al comments. We thank Andrew Stewart and Carl Struthers, Te Papa Well�ngton, for supply of Squalus t�ssue samples from New Zealand. Peter Sm�th was supported by a FRST contract on the molecular taxonomy of NZ fishes. This work was also supported (�n part) by the NZ Foundat�on for Research Sc�ence and Technology through the Te Papa B�osystemat�cs of NZ EEZ F�shes subcontract w�th�n NIWA’s mar�ne B�od�vers�ty and B�osecur�ty OBI programme (contract C01X0502).

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122

Cluster A

Cluster B

Cluster C

Figure 1. K2P d�stance ne�ghbour jo�n�ng tree for the 16 spec�es of Squalus show�ng the three major clusters of spec�es referred to �n text and F�gs 2–4. K2P d�stance bar g�ven.


Figure 2. K2P d�stance ne�ghbour jo�n�ng tree for Squalus sequences in Cluster A. In this and figs 3–7, species, sample designation and country of origin are shown. T fin = Taiwan highfin, L hfin = Lombok highfin. Bootstrap values > 50% shown. K2P d�stance bar g�ven.

124

Figure 3. K2P d�stance ne�ghbour jo�n�ng tree for Squalus sequences �n Cluster B. Bootstrap values > 50% shown. K2P d�stance bar g�ven.

Figure 4. K2P d�stance ne�ghbour jo�n�ng tree for Squalus sequences �n Cluster C. Bootstrap values > 50% shown. K2P d�stance bar g�ven.


Figure 5. K2P d�stance UPGMA tree for Squalus sequences in Cluster A. T hfin = Taiwan highfin, L hfin = Lombok highfin. Bootstrap values > 50% shown. K2P d�stance ax�s g�ven.

126

Figure 7. K2P d�stance UPGMA tree for Squalus sequences �n Cluster C. Bootstrap values > 50% shown. K2P d�stance ax�s g�ven.

Figure 6. K2P d�stance UPGMA tree for Squalus sequences �n Cluster B. Bootstrap values > 50% shown. K2P d�stance ax�s g�ven.


Table 1. Summary deta�ls of the 127 spec�mens of Squalus sequenced for the barcode reg�on of mtDNA COI.CSIRO = Austral�an Nat�onal F�sh Collect�on, CSIRO, Hobart, Austral�a; KAUM = Kagosh�ma Un�vers�ty Museum, Kagosh�ma, Japan; MLML = Moss Land�ng Mar�ne Laboratory, Cal�forn�a, USA; MZB = Museum Zoolog�cum Bogor�ense, Jakarta, Indones�a; NMNZ = Nat�onal F�sh Collect�on at the Museum of New Zealand, Te Papa Well�ngton, New Zealand; UF = Un�vers�ty of Flor�da, Flor�da, USA.

Species Sample ID number

GenBank No.

Voucher No. Voucher location Collection site

S. acanthias 16 S – not reta�ned – USA, Wash�ngton coastS. acanthias 17 S – not reta�ned – USA, Wash�ngton coastS. acanthias BW–A2327 – not reta�ned – USA, Puget SoundS. acanthias BW–A2328 – not reta�ned – USA, Puget SoundS. acanthias BW–A2329 – not reta�ned – USA, Puget SoundS. acanthias BW–A2330 – not reta�ned – USA, Puget SoundS. acanthias BW–A2331 – not reta�ned – USA, Puget SoundS. acanthias BW–A2332 – not reta�ned – Japan, off Hokka�do IslandS. acanthias BW–A2333 – not reta�ned – Japan, off Hokka�do IslandS. acanthias BW–A2334 – not reta�ned – Japan, off Hokka�do Island

S. acanthias 42P40172 – NMNZ P 40172 Well�ngton, NZ New Zealand, North IslandS. acanthias BW–A083 DQ108279 not reta�ned – Austral�a, Tasman�aS. acanthias BW–A084 DQ108280 not reta�ned – Austral�a, Tasman�aS. acanthias BW–A085 DQ108281 not reta�ned – Austral�a, Tasman�aS. acanthias BW–A086 DQ108282 not reta�ned – Austral�a, Tasman�aS. acanthias BW–A087 DQ108267 CSIRO H 4876–01 CSIRO, Hobart Austral�a, Tasman�aS. acanthias BW–A2312 – not reta�ned – USA, Gulf of Ma�neS. acanthias BW–A2313 – not reta�ned – USA, Gulf of Ma�neS. acanthias BW–A2314 – not reta�ned – USA, Gulf of Ma�neS. acanthias BW–A2315 – not reta�ned – USA, Gulf of Ma�neS. acanthias BW–A2316 – not reta�ned – USA, Gulf of Ma�neS. acanthias BW–A2317 – not reta�ned – Ch�le, Isla MochaS. acanthias BW–A2318 – not reta�ned – Ch�le, Isla MochaS. acanthias BW–A2319 – not reta�ned – Ch�le, Isla MochaS. acanthias BW–A2320 – not reta�ned – Ch�le, Isla MochaS. acanthias BW–A2321 – not reta�ned – Ch�le, Isla MochaS. acanthias BW–A2322 – not reta�ned – UK, Celt�c SeaS. acanthias BW–A2323 – not reta�ned – UK, Celt�c SeaS. acanthias BW–A2324 – not reta�ned – UK, Celt�c SeaS. acanthias BW–A2325 – not reta�ned – UK, Celt�c SeaS. acanthias GenBank Y18134 – – Iceland, Faxaflói

S. albifrons BW–A030 – CSIRO H 3589–01 CSIRO, Hobart Austral�a, New South WalesS. albifrons BW–A107 DQ108254 CSIRO H 4627–01 CSIRO, Hobart Austral�a, New South WalesS. albifrons BW–A108 DQ108255 CSIRO H 4704–01 CSIRO, Hobart Austral�a, New South WalesS. albifrons BW–A109 DQ108256 CSIRO H 4704–02 CSIRO, Hobart Austral�a, New South WalesS. albifrons BW–A110 DQ108257 CSIRO H 4705–01 CSIRO, Hobart Austral�a, New South Wales

S. brevirostris BW–A2284 – CSIRO H 6293–29 CSIRO, Hobart Ta�wan

S. brevirostris BW–A2285 – CSIRO H 6293–30 CSIRO, Hobart Ta�wanS. brevirostris BW–A2286 – CSIRO H 6293–31 CSIRO, Hobart Ta�wanS. brevirostris BW–A2990 – KAUM I 184 KAUM, Kagosh�ma Japan, East Ch�na Sea

128


GenBank No

Voucher No Voucher location Collection site

S. brevirostris BW–A2991 – KAUM I 185 KAUM, Kagosh�ma Japan, East Ch�na SeaS. brevirostris BW–A2992 – KAUM I 186 KAUM, Kagosh�ma Japan, East Ch�na SeaS. brevirostris BW–A2993 – KAUM I 187 KAUM, Kagosh�ma Japan, East Ch�na SeaS. brevirostris BW–A2994 – KAUM I 377 KAUM, Kagosh�ma Japan, East Ch�na Sea

S. chloroculus BW–A100 DQ108263 not reta�ned – Austral�a, South Austral�aS. chloroculus BW–A101 DQ108264 CSIRO H 4775–01 CSIRO, Hobart Austral�a, V�ctor�aS. chloroculus BW–A2214 – CSIRO H 5941–01 CSIRO, Hobart Austral�a, Tasman�aS. chloroculus BW–A097 DQ108260 not reta�ned – Austral�a, South Austral�aS. chloroculus BW–A098 DQ108261 not reta�ned – Austral�a, South Austral�aS. chloroculus BW–A099 DQ108262 not reta�ned – Austral�a, South Austral�aS. chloroculus P38327 – NMNZ P 38327 Well�ngton, NZ Tasman Sea, Norfolk R�dge

S. crassispinus BW–A116 DQ108247 CSIRO H 4649–03 CSIRO, Hobart Austral�a, Western Austral�aS. crassispinus BW–A117 DQ108248 CSIRO H 4649–04 CSIRO, Hobart Austral�a, Western Austral�a

S. edmundsi BW–A102 DQ108265 CSIRO H 2608–16 CSIRO, Hobart Austral�a, Western Austral�aS. edmundsi BW–A103 DQ108266 CSIRO H 2605–05 CSIRO, Hobart Austral�a, Western Austral�aS. edmundsi BW–A104 DQ108251 CSIRO H 2605–06 CSIRO, Hobart Austral�a, Western Austral�aS. edmundsi BW–A105 DQ108252 CSIRO H 2605–07 CSIRO, Hobart Austral�a, Western Austral�aS. edmundsi BW–A111 DQ108258 CSIRO H 3969–15 CSIRO, Hobart Austral�a, Western Austral�aS. edmundsi BW–A112 DQ108243 not reta�ned – Austral�a, Western Austral�aS. edmundsi BW–A113 DQ108244 not reta�ned – Austral�a, Western Austral�aS. edmundsi BW–A114 DQ108245 not reta�ned – ?S. edmundsi BW–A115 DQ108246 not reta�ned – ?S. edmundsi BW–A2287 – CSIRO H 5857–09 CSIRO, Hobart Indones�a, Bal�S. edmundsi BW–A2288 – CSIRO H 5857–10 CSIRO, Hobart Indones�a, Bal�S. edmundsi BW–A2290 – CSIRO H 5875–05 CSIRO, Hobart Indones�a, LombokS. edmundsi BW–A2291 – CSIRO H 5875–06 CSIRO, Hobart Indones�a, LombokS. edmundsi BW–A2600 – not reta�ned – Indones�a, Lombok

S. grahami BW–A120 DQ108235 CSIRO H 4682–01 CSIRO, Hobart Austral�a, New South WalesS. grahami BW–A121 DQ108236 CSIRO H 4682–02 CSIRO, Hobart Austral�a, New South WalesS. grahami BW–A122 DQ108237 CSIRO H 4682–03 CSIRO, Hobart Austral�a, New South WalesS. grahami BW–A123 DQ108238 CSIRO H 4682–04 CSIRO, Hobart Austral�a, New South WalesS. grahami BW–A124 DQ108239 CSIRO H 4623–03 CSIRO, Hobart Austral�a, New South WalesS. grahami BW–A94 DQ108273 CSIRO H 4623–02 CSIRO, Hobart Austral�a, New South Wales

S. griffini 10P41775 – NMNZ P 41775 Well�ngton, NZ New Zealand, South IslandS. griffini 19P40888 – NMNZ P 40888 Well�ngton, NZ New Zealand, Raoul IslandS. griffini 21P40347 – NMNZ P 40347,

TS1029Well�ngton, NZ New Zealand, North Island

S. griffini 25P40346 – NMNZ P 40346 Well�ngton, NZ New Zealand, North IslandS. griffini 26P40347 – NMNZ P 40346,

TS1027Well�ngton, NZ New Zealand, North Island

S. griffini 27P41774 – NMNZ P 41774,765TL

Well�ngton, NZ New Zealand, South Island

Table 1. cont’d.



GenBank No


S. griffini 28P41774 – NMNZ P 41774,TL690

Well�ngton, NZ New Zealand, South Island

S. griffini 2P35352 – NMNZ P 35352 Well�ngton, NZ New Zealand, Kermadec R�dgeS. griffini 30P41196 – NMNZ P 41196 Well�ngton, NZ New Zealand, Chatham R�seS. griffini 32P1193 – NMNZ P 41993 Well�ngton, NZ New Zealand, Chatham R�seS. griffini 4P35270 – NMNZ P 35270 Well�ngton, NZ New Zealand, Chatham R�seS. griffini 7P40346 – NMNZ P 40346,

826TLWell�ngton, NZ New Zealand, North Island

S. griffini 8P4034 – NMNZ P 40346,807TL

Well�ngton, NZ New Zealand, North Island

S. griffini P042491 – NMNZ P 42491,TS1732

Well�ngton, NZ New Zealand, Challenger Plateau

S. griffini P042492 – NMNZ P 042492,TS1733

Well�ngton, NZ New Zealand, Challenger Plateau

S. hemipinnis BW–A2302 – CSIRO H 5631–02 CSIRO, Hobart Indones�a, Bal�S. hemipinnis BW–A2303 – CSIRO H 5631–01 CSIRO, Hobart Indones�a, Bal�S. hemipinnis BW–A2304 – CSIRO H 5631–05 CSIRO, Hobart Indones�a, Bal�S. hemipinnis BW–A2305 – CSIRO H 5631–03 CSIRO, Hobart Indones�a, Bal�S. hemipinnis BW–A2306 – CSIRO H 5631–04 CSIRO, Hobart Indones�a, Bal�S. hemipinnis BW–A2601 – not reta�ned – Indones�a, Lombok

S. japonicus BW–A2292 – CSIRO H 6294–26 CSIRO, Hobart Ta�wanS. japonicus BW–A2293 – CSIRO H 6294–27 CSIRO, Hobart Ta�wanS. japonicus BW–A2294 – CSIRO H 6294–31 CSIRO, Hobart Ta�wanS. japonicus BW–A2295 – UF 148938 UF, Flor�da Ta�wanS. japonicus BW–A2296 – TAI–119 MLML, Cal�forn�a Ta�wan

S. megalops BW–A088 DQ108268 not reta�ned – Austral�a, Bass Stra�tS. megalops BW–A089 – not reta�ned – Austral�a, Bass Stra�tS. megalops BW–A090 DQ108269 not reta�ned – Austral�a, Bass Stra�tS. megalops BW–A091 DQ108270 not reta�ned – Austral�a, Bass Stra�tS. megalops BW–A092 DQ108271 CSIRO H 3762–01 CSIRO, Hobart Austral�a, Bass Stra�t

S. montalbani BW–A106 DQ108253 CSIRO H 2606–05 CSIRO, Hobart Austral�a, Western Austral�aS. montalbani BW–A2298 – CSIRO H 5857–17 CSIRO, Hobart Indones�a, Bal�S. montalbani BW–A2299 – CSIRO H 5857–18 CSIRO, Hobart Indones�a, Bal�S. montalbani BW–A2300 – CSIRO H 5857–06 CSIRO, Hobart Indones�a, JavaS. montalbani BW–A2301 – MZB 15022 MZB, Jakarta Indones�a, JavaS. montalbani BW–A093 DQ108272 CSIRO H 4623–01 CSIRO, Hobart Austral�a, New South WalesS. montalbani BW–A095 DQ108274 CSIRO H 4623–04 CSIRO, Hobart Austral�a, New South WalesS. montalbani BW–A096 DQ108259 CSIRO H 4623–05 CSIRO, Hobart Austral�a, New South WalesS. montalbani P042490 – NMNZ P 42490,

TS1731Well�ngton, NZ New Zealand, Challenger

Plateau

S. nasutus BW–A118 DQ108249 not reta�ned – Austral�a ?S. nasutus BW–A119 DQ108250 not reta�ned – Austral�a ?S. nasutus BW–A2199 – not reta�ned – Indones�a, Lombok

Table 1. cont’d.

130


GenBank No


S. nasutus BW–A2307 – MZB 15008 MZB, Jakarta Indones�a, Bal�S. nasutus BW–A2308 – MZB 15009 MZB, Jakarta Indones�a, Bal�S. nasutus BW–A2309 – CSIRO H 5860–01 CSIRO, Hobart Indones�a, JavaS. nasutus BW–A2310 – CSIRO H 5680–03 CSIRO, Hobart Indones�a, JavaS. nasutus BW–A2311 – CSIRO H 5680–02 CSIRO, Hobart Indones�a, Java

S. raoulensis 18P41678 – NMNZ P 41678 Well�ngton, NZ New Zealand, Raoul IslandS. raoulensis 20P41678 – NMNZ P 42572 Well�ngton, NZ New Zealand, Raoul IslandS. raoulensis 5P34436 – NMNZ P 34436 Well�ngton, NZ New Zealand, Raoul Island

S. sp. (Lombok highfin)

BW–A2289 – CSIRO H 5788–06 CSIRO, Hobart Indones�a, Lombok

S. sp. (Ta�wan highfin)

BW–A2282 – TAI–001 Ta�wan? Ta�wan

S. sp. (Ta�wan highfin)

BW–A2283 – CSIRO H 6292–10 CSIRO, Hobart Ta�wan

Table 1. cont’d.

descriptions of new dogfishes · in the follow ng ser es of papers, a rap d taxonom c approach s...

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